(Oceanography and Marine Biology - An Annual Review) Alan Ansell - Oceanography and Marine Biology - An Annual Review, Volume 34-CRC Press (1996)

Cover
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OCEANOGRAPHY and MARINE BIOLOGY
AN ANNUAL REVIEW
Volume 34
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OCEANOGRAPHY and MARINE BIOLOGY

AN ANNUAL REVIEW
Volume 34
Editors
A.D.Ansell
R.N.Gibson
Margaret Barnes
The Dunstaffnage Marine Laboratory Oban, Argyll, Scotland

Founded by Harold Barnes
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© A.D.Ansell, R.N.Gibson and Margaret Barnes, 1996

This book is copyright under the Berne Convention.
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First published in 1996 by UCL Press
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ISBN 0203501268 Master ebook ISBN
ISBN 0203231325 (OEB Format)

ISBN:
1857285816 (Print Edition) HB
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CONTENTS
Preface vii
Physical oceanography of the Bay of Bengal and Andaman Sea 1

M.J.Varkey, V.S.N.Murty & A.Suryanarayana
Nutrient fluxes to planktonic osmotrophs in the presence of fluid motion 71
L.KarpBoss, E.Boss & P.A.Jumars
Factors affecting grazerepiphyte interactions in temperate seagrass meadows 109
P.Jernakoff, A.Brearley & J.Nielsen
Beach clam fisheries 163
Anton McLachlan, Jenifer E.Dugan, Omar Defeo, Alan D.Ansell, David M.Hubbard, Eduardo Jaramillo & Pablo E.Penchaszadeh
Biology of the gastropod family Littorinidae. I. Evolutionary aspects 233
Christopher D.McQuaid
Biology of the gastropod family Littorinidae. II. Role in the ecology of intertidal and shallow marine ecosystems 263
Christopher D.McQuaid
Pedunculate cirripedes of the genus Pollicipes 303
Margaret Barnes
Coastal fisheries in the Pacific islands 395
P.Dalzell, T.J.H.Adams & N.V.C.Polunin
Author index 533

Systematic index 557
Subject index 567
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Page vii
PREFACE
The thirtyfourth volume of this series of annual reviews contains eight articles that cover, as usual, a wide range of topics from physical oceanography through biology
and ecology to fisheries. This year there are contributors from ten countries from Europe, Africa, the Americas, Asia and Australia, and it is gratifying to the editors that
the series continues to attract such a wide support among the international marine science community. Our editorial policy continues to be that of maintaining a high
standard of authoritative review, both by soliciting articles in subjects where we perceive that a comprehensive coverage would be timely, and by accepting suitable
reviews that are offered to us. We welcome suggestions of subjects that might be reviewed in future volumes. Since an annual publication schedule necessarily places
some constraints on the timetable for submission and acceptance of manuscripts, however, we advise potential contributors to make contact at an early stage of
preparation, so that they may time their submission to minimize the delay before publication. Our publishers, UCL Press, now routinely use authors’ disks in the initial
preparation of text, tables and, in some cases, figures, hopefully helping to minimize errors in production. We are grateful to them for continuing to maintain the annual
appearance of these reviews and to all our contributors for their patience and cooperation in dealing with editorial queries and requests.
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Page 1
PHYSICAL OCEANOGRAPHY OF THE BAY OF BENGAL AND

ANDAMAN SEA
M.J.VARKEY, V.S.N.MURTY & A.SURYANARAYANA
National Institute of Oceanography, Dona Paula, 403004 India
Oceanography and Marine Biology: an Annual Review 1996, 34, 1–70

© A.D.Ansell, R.N.Gibson and Margaret Barnes, Editors UCL Press
Abstract Physical oceanography of the Bay of Bengal and Andaman Sea is reviewed for the first time. All available information for over 50 years is
consolidated in this review. To begin with, information on peripheral or related aspects of climate of the hinterland, such as pressure, rainfall, storms,
winds, sunshine etc., are presented. The Bay of Bengal and Andaman Sea are divided into two regions, viz., the coastal belt and the open ocean
areas, on the basis of large differences in water properties. Considerable variation in salinity and density exists all over the coastal areas due to
abundant freshwater discharges, especially during the rainy months. The coastal circulation is mainly driven by river runoff and wind thrust and is
influenced by branches of the Equatorial Current system. Strong winddriven upwelling and sinking are seen along the eastern Indian coast. The
surface circulation system in the Bay of Bengal and Andaman Sea can be divided into three gyres, viz., (a) the northwestern gyre, (b) the large gyre in
the southern Bay of Bengal and (c) the Andaman gyre. Gyres are noted also at 500m and 1000m. The net water exchanges [Evaporation (E)+
Precipitation (P)+Runoff (R)] at surface, for the whole area north of 5°N are −11.5cm and +63.7cm during the winter monsoon and summer
monsoon, respectively. The net water transports across 6°N between surface and 1250m, are found to be −79×1012m3 and +129×1012m3 for winter
monsoon and summer monsoon periods, respectively. High salinity water masses from the Red Sea, Persian Gulf and eastern and southern Arabian
Sea flow into the Bay of Bengal and play an important role in its water structure. A strong gradient of surface temperature is observed during winter
(25.5 to 28.5°C) from the head of the Bay to 5°N; similarly for salinity (21.0 to 34.5ppt) during summer. Variation in mixed layer thickness is greater
during summer (about 25 to 100m) and less during the winter (60 to 100m). Abyssal water flows and properties are least studied in the Bay of
Bengal.
Introduction
The Indian Ocean was poorly studied before the International Indian Ocean Expedition (IIOE), which was established to bridge this information gap. This international
endeavour, spread mainly over the period 1962–66, produced substantial data that were later analyzed and published by the international community of marine
scientists. These papers, monographs, books and atlases dramatically increased knowledge of the Indian Ocean. Considering the Bay of Bengal (Fig. 1) alone, prior to
the IIOE the situation was very bad (Currie 1996) and the improvement after IIOE has been tremendous. During the last two decades since the IIOE, a considerable
amount of scientific work has been carried out in the Bay of Bengal and Andaman Sea by many organizations of various nations, especially India, and more data and
information have been added. Here the authors aim at consolidating the scattered pieces of information present in various papers, reports and books into a systematic
and concrete form.
For the ensuing discussion, the Bay of Bengal and Andaman Sea together are defined as the oceanic area north of 5°N, excluding the Malacca Strait and Gulf of
Mannar, bordered by the Indian subcontinent, Burma, Thailand, Malay Peninsula and Sumatra (hereafter, for
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convenience, termed as “the Bay” when referred to together). The Malacca Strait and Gulf of Mannar are excluded because of their hydrographic insignificance
compared with the large and deep Bay. Viewing the Bay in a hemispherical perspective, with emphasis on the North Equatorial Current of the eastern Indian Ocean, it
can be clearly seen (Figs 23a, 42, see pp.32 and 59) that 5°N rightly forms the southern boundary line of the Bay between Sri Lanka and Sumatra, both physically and
dynamically. The northern boundary around 5°N of the North Equatorial Current, which reverses seasonally, makes the southern dynamic boundary of the Bay with
meanders extending into the Bay north of 5°N (Anonymous 1960). The Andaman Sea is almost a separate sea and forms the eastern part of the northeastern Indian
Ocean, partitioned from the western Bay by the AndamanNicobar island chain. These two parts of the northeastern Indian Ocean are connected deeply in the
Preparis Channel, Ten Degree Channel, and the Great Channel (Fig. 1). The study area has a surface area of 2.8×106km2 and covers about 6% of the surface of the
Indian Ocean. Water depths vary
Figure 1 Map of Bay of Bengal and surrounding oceanic and land areas.
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from 4000m around 5°N to 200m at the shelf edge. The continental shelf along the east coast of India is very narrow (<45km), but the shelf areas off the mouths of the
Ganges, Irrawady and Salween are very wide (>200km).
As defined above, the Bay forms the northeastern part of the Indian Ocean. The Arabian Sea, the northwestern part, is completely separated from the Bay by the
peninsula of India. The Indian Ocean, as a whole, is characterized by the reversing monsoons; summer monsoon during June, July, August and September and winter
monsoon during November, December, January and February (peak months only). The North Equatorial Current in the Indian Ocean completely reverses and changes
markedly in response to the reversing winds. The conspicuousness of this phenomenon is so striking that this Ocean is considered as the best oceanic region for study
of the response of ocean to atmosphere. The tropospheric north Indian Ocean is characterized by the presence of high salinity water masses from the Persian Gulf, Gulf
of Aden and northern Arabian Sea. The study area is very interesting for its low salinity surface water caused by large river runoff from the Indian subcontinent and
Burma.
Observations in the Bay started in the 19th century with expeditions of the NOVARA (1857–59) and VALDIVIA (1898–99). The data collected during the cruises
of RMIS INVESTIGATOR were analyzed and documented by Sewell in various reports of Memoirs of the Asiatic Society of Bengal (Sewell 1925, 1928, 1929,
1932). During the 1950s many small surveys were carried out by Andhra University, limited to the western coastal areas of the Bay. Throughout the duration of the
IIOE many cruises were conducted in the Bay during various months and the availability of data for winter, especially for deep waters, increased tremendously. The
voluminous data collected for the Indian Ocean during the IIOE were analyzed and presented in the form of an atlas by Wyrtki (1971) and this atlas remains as the
single authoritative reference work for the oceanography of the Indian Ocean. With the commissioning of the RV GAVESHANI, the first research vessel of India, by
the National Institute of Oceanography in 1976, another phase of research started in the Indian Ocean. Since then the availability of data has greatly increased
especially for the summer monsoon and transition periods. For the transition seasons, however, data coverage is still poor, as can be seen from the distribution charts.
The oceanography of the Bay and coastal regions around the Indian subcontinent is now known in much more detail.
This review is divided into 5 sections: (1) Atmospheric and land environment (2) Coastal oceanography (3) Deep Bay oceanography (4) Currents and water
transports and (5) Summary. Section 1 provides background knowledge, and emphasizes seasonal aspects of runoff, evaporation, precipitation, etc. Section 2 deals
specifically with shallow coastal areas becuase of large variations in the properties from the open Bay. Section 3 covers the open, deep Bay, which shows large water
mass inhomogeneities caused by the flow of high salinity water from the Arabian Sea and the penetration of the seasonally reversing North Equatorial Current across 5°
N. Section 4 deals exclusively with currents in a detailed way, bringing out some clear patterns. In sections 2, 3 and 4, the historical literature is initially presented in
three groups: (1) prior to IIOE, (2) during and after IIOE and (3) after the commissioning of RV GAVESHANI. After the historical literature, all results are analyzed
and interpreted with the help of typical diagrams mainly from the author’s own work, but with some reproduced from other works. For the discussions in sections 2, 3
and 4, the seasons are grouped as follows: (a) presummer monsoon (AprilMay), (b) summer monsoon (JuneSeptember), (c) postsummer monsoon (Octobermid
November) and (d) winter monsoon (mid NovemberMarch). Here it should be mentioned that the pre and postsummer monsoons are regarded as transition
periods.
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Atmospheric and land environments
General climatic aspects

For a good review of the physical oceanography of the Bay it is necessary to understand the climatological features of the study area and its neighbourhood. The
hinterland of the Bay is defined here as the extensive land area that contains the tributaries and the distributaries of the major rivers which flow into the Bay. This
hinterland acquires special importance for this study because of the extensive river runoff into the Bay and its effects on water properties. Thus, the catchment areas, the
plains and the deltas of the big rivers—Brahmaputra, Cauvery, Damodar, Ganges, Godavari, Irrawady, Krishna, Mahanadi, Mahaweli, Pennar, and Salween—fall into
the hinterland (Fig. 1). This section on climatology and related aspects is of peripheral interest only and all the information is extracted from published works.
The Bay and its hinterland cover a wide range of climatic features. Since the climates of the study area are primarily influenced by the monsoons, a climatic
classification based on amounts of precipitation, particularly on the lengths of dry and wet seasons, is relevant here. Such a classification for the area is presented by
Landsberg et al. (1966) and the following information is extracted from it. The hinterland of the Ganges, Brahmaputra, Damodar, Mahanadi and Mahaweli (Figs 1, 2)
experience tropical rainy climates and tropical humid
Figure 2 Climatic classification of Bay of Bengal and hinterland (reproduced from Landsberg et al. 1966).
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summer climates with humid winters, and is grouped under V1 (rain forest climates) and V2 (moist savanna with 5–7.5 months dry season). The Irrawady and Salween
in Burma, cover climatic scales VI, V2 and V3 (dry savanna with 5–7.5 months dry season). The rivers that flow into the Bay across the southeast coast of India are
associated with land masses of climatic group V3 and V4 (tropical semideserts of dry climates with humid winters). The oceanic area that lies between the Burma
Thailand coastline and the AndamanNicobar Island chain comes under V1. Most of the Bay area, west of the Andamans, is grouped under V2 except for a narrow
belt off the southeast coast of India which is placed under V3.
Total solar radiation (direct solar radiation+diffuse sky radiation) varies from 120 to 180kcalcm2yr−1 in the whole area (Bay+hinterland) (Landsberg et al. 1966).
Representative mean sunshine (h/July) for the summer period is 100 to 150h, whereas for winter (h/January) it is 200 to 250h. The annual mean sunshine duration for
the study area is 1600 to 3000h (Landsberg et al. 1966). Mean cloudiness (in tenths) in the Bay for January is between 3 and 4 whereas for July the range is 3 to 8
(Ramage 1984).
Mean sea level pressure decreases from 1016mb over the Tibetan Plateau to 1010mb at the equator in January (Rao, Y.P. 1981, Ramage 1984). In July the
pressure decreases from 1010mb at the Equator to 1000mb over Tibet (Ramage 1984, Rao, Y.P. 1981). The change in pressure gradients between the winter
monsoon and summer monsoon is very marked. The Bay experiences monthly mean winds mainly from the northeast in the range of 6 to 12 knots during December
(Fig. 3). In the Andaman Sea the directions are more easterly and at the head of the Bay more northerly. During April (Fig. 3), wind directions change from southerly
near Sri Lanka to northerly over the coast of Burma, turning in a big anticyclonic pattern centred on the Ten Degree Channel. Wind speeds are less than 8 knots over
much of the Bay. During July, a typical summer month, wind direction is mainly from the southwest (Fig. 4) with speeds ranging from 10 to 18 knots. During October
(Fig. 4), a postsummer monsoon transition month, wind directions vary greatly all over the Bay and speeds are less than 4 knots. Wind steadiness over the Bay during
January and July, typical monsoon months, is very high (>90%) (Hastenrath & Lamb 1979). During October, steadiness is very low (c.30%) and in April high (c.80%)
in the northwestern Bay and low (c.30%) in the Andaman
Figure 3 Wind speeds and directions during December and April for the Bay of Bengal (reproduced from Hastenrath and Lamb 1979).
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Figure 4 Wind speeds and directions during July and October for the Bay of Bengal (reproduced from Hastenrath & Lamb 1979).
Sea and southern Bay (Hastenrath & Lamb 1979). Over the Indian subcontinent mean winds of 0–5 knots with varying directions prevail during January and July (Rao,
Y.P. 1981). Over Malaysia, Thailand and Burma also, winds are northeasterly during January, a fairly typical winter month, and are southwesterly during July, a
summer month (Nieuwolt 1981).
Tropical and subtropical cyclones account for stormy winds and heavy rainfalls over the northern Indian Ocean and surrounding areas. Over the Bay storms and
depressions are observed mostly from June to November whereas in the Arabian Sea a bimodal distribution, with MayJuly and OctoberNovember as the two
peaks, is observed (Rao, K.N. 1981). Even though the storms and depressions are spread over the complete area between 5°N to 28°N and 55°E to 98°E, they
concentrate at the head of the Bay and Arabian Sea (Rao, K. N. 1981). Warmcore monsoon depressions are the prime rain producer over the northern Bay (Ramage
1984). Tropical cyclones in the Bay during the postmonsoon transition period are associated with very heavy rains and stormy winds especially in the coastal areas
(Rao, K.N. 1981).
Details of mean annual rainfall on global and regional bases are presented by Baumgartner & Reichel (1975) and the following details on rainfall are extracted from
that work. The eastern half of Sri Lanka experiences rains in the range of 140–250cmyr−1. The west coasts of Burma, Thailand, Malaysia and northern Sumatra
experience heavy rainfalls in the range 200–400cmyr−1 while the interior of Burma (Shan Plateau) receives only 100–200cmyr−1. Rainfall increases from 60cmyr−1
over the southeast coast of India to 200cmyr−1 over Bangladesh. The Gangetic Plains and the subHimalayan ranges also receive heavy rains (200–400cmyr−1). Rains
over the Bay and hinterland show strong seasonality. The southeast coast of India has a winter rainfall maximum (Ramage 1984). The rest of the east coast of India,
Bangladesh and Burma have a summer rainfall maximum (Ramage 1984). Sri Lanka, the western tip of Sumatra, Malaysia and Thailand have a bimodal pattern with
maxima during summer and winter (Ramage 1984).
The following details on evaporation and discharge are also extracted from Baumgartner & Reichel (1975). Mean annual evaporation rates over the eastern half of
Sri Lanka, the northern tip of Sumatra, the western coastal areas of Malaysia, Thailand and Burma and the
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Gangetic Delta are in the range of 110–140cmyr−1. Interior Burma (Shan Plateau) experiences evaporation rates within the range 80–100cmyr−1. Over the sub
Himalayan ranges and Gangetic plains evaporation values range between 40cmyr−1 and 110cmyr−1. Evaporation rates over eastern India are within the range 60–
100cmyr−1. Discharges (precipitation minus evaporation) over eastern Sri Lanka vary between 20cmyr−1 and 100cm yr−1. Over the northern tip of Sumatra and
coastal areas of Burma, discharge rates range between 100cmyr−1 and 200cmyr−1. Interior Burma (Shan Plateau, has rates of 20 to 60cmyr−1. The eastern central belt
of India has rates between 5cmyr−1 and 40cmyr−1. Discharge rates of the subHimalayan ranges are very high (100–300cmyr−1).
Runoffs of major rivers

The hinterland has a number of rivers emptying into the Bay. Details on flow rates and other aspects of the major rivers are obtained from UNESCO (1969, 1971a,b),
Rao (1979) and Anonymous (1981a,b). Some of the small rivers (e.g. Palar in India) have only seasonal or negligible flows and are not considered in the ensuing
discussion. For each river the flow ratios for WM (winter monsoon including December, January and February only) and SM (summer monsoon including June, July
and August only) are computed as the ratio of the total seasonal runoff to total annual runoff (Table 1). For cases where the monthly distributions of the runoffs were
not available the mean annual runoff values were distributed monthwise considering the monthly distributions of other river or rivers that flow over areas with similar
climatic conditions (see Fig. 2); for example, between the Arakan ranges in Burma and subHimalayan ranges and between the Shan Plateau in Burma and southeast
central India. The runoff values for the Ganges and Brahmaputra are for Farakka point and the Indian boundary, respectively, far from the Bay. Hence for the Meghna
area in Bangladesh, the deltaic areas of the Ganges and Brahmaputra, the discharge rate is computed as the product of a spatial mean discharge height of 91cm
obtained from Baumgartner & Reichel (1975)
Table 1 Annual and seasonal runoffs of major rivers flowing into the Bay
(WM=Dec.+Jan.+Feb., SM=June+July+Aug.).
Annual mean (m3 s−1) Total run off volume (×109m3)

River WM ratio SM ratio WM SM
Mahaweli 226 0.430 0.163 31.065 1.159
Godavari 3180 0.022 0.554 2.204 55.565
Krishna 1730 0.021 0.619 1.140 33.798
Mahanadi 1710 0.021 0.599 1.137 32.320
Pennar 95 0.203 0.138 0.607 0.414
Damodar 329 0.031 0.561 0.317 5.823
Ganges 11892 0.065 0.493 24.497 184.736
Brahmaputra 16186 0.059 0.496 30.367 252.934
Cauvery 664 0.203a
0.138 a 4.245 2.893
Irrawady 13018 0.049b 0.529b 19.966 217.235
Salween 5421 0.049b 0.529b 8.314 90.462
Meghna (in Bangladesh) 4215 0.059c
6.496 c 7.908 65.867
Total 103.767 943.206
a. Same as that of Pennar
b. Mean value from ratios of Mahanadi, Ganges and Brahmaputra
c. Same as that of Brahmaputra
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for the Bangladesh area and the surface area of Bangladesh (about 1.5×105km2; see Varkey 1986 for more details). Total surface area of the Bay is worked out as
2.8×106km2. Hence using the grand total runoff value (Table 1), the runoff heights for the Bay are 3.7cm (103.8×105hectare metres) and 34cm (943.2×105hectare
metres) for WM and SM, respectively. It is estimated from data presented in Table 1 and Anonymous (1988) that the Bay receives about 2125km3 of runoff yearly
from the east coast of India, Burma and Bangladesh.
Evaporation and precipitation in the Bay

In the Indian Ocean mean annual evaporation shows two maxima; one (>70cm) between 10°–20°S and 70°–90°E, and the other (>130cm) off the Somali coast
between 0–10°N and 45°–65°E (Venkateswaran 1956, Baumgartner & Reichel 1975). These two maxima are observed throughout the year except during
SeptemberNovember when the northwestern maximum subsides to a minimum (<20cm/quarter). Throughout the Bay yearly mean values (100–1 10cmyr−1), WM
mean values (c.30cm/quarter) and SM mean values (30–35 cm/quarter) remain almost uniform (Venkateswaran 1956). Comparatively, the Arabian Sea experiences
more evaporation in all seasons except in the postmonsoon period of SeptemberNovember (Venkateswaran 1956). Table 2 presents the seasonal evaporation rates
for WM and SM for the different zones in the Bay. These values are estimated using data on monthly heat loss presented in the IIOE atlas (Ramage et al. 1972). The
values presented in Table 2 agree very well with those in Venkateswaran (1956).
Distribution of components of the mean water balance equation [P=E+D (discharge)] for the global Earth, for regional land masses and oceans are presented in
Baumgartner & Reichel (1975) in the form of maps of longterm average contours of P, E and D. Annual mean precipitation values presented in Table 3 are extracted
from the above work. The seasonal indices (Table 3) for calculating seasonal precipitation are worked out from rainfall data published in Schutz (1967), Anonymous
(1971), Walter et al. (1975) and Rao (1981) for the Indian subcontinent, South East Asia and Bay Islands, as the fraction of the total seasonal (WM and SM)
precipitation to the total yearly precipitation (see Varkey 1986 for more details).
Table 2 Evaporation (cm/season) over the Bay for winter and summer. Summer values are shown in brackets.
Latitude belt (°N) Longitudinal belt (°E)

80–85 85–90 90–95 95–100 Mean
>20 _ 34.25 28.44 _ 31.34
(26.78) (30.46) (28.62)
15–20 36.51 31.58 26.48 22.57 31.52
(34.89) (30.45) (34.09) (27.39) (31.71)
10–15 37.32 35.61 29.27 33.15 33.84
(36.77) (40.16) (33.33) (30.10) (35.09)
5–10 38.86 39.41 37.60 39.87 38.94
(39.95) (41.21) (43.88) (34.83) (39.94)
Mean 37.56 35.21 30.45 31.86
(37.20) (34.63) (35.44) (30.77)
Grand Mean=33.77
(34.51)
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Net estimates at surface

Using the values presented in Tables 1, 2 and 3, net water exchanges at the surface of the Bay for the different zones are worked out and the results are presented in
Table 4. Here, evaporation is considered to be negative since water is removed from the surface and precipitation and runoff are taken to be positive since water is
added to the surface. Net volumes are computed using the surface areas of the various zones. Here it is assumed that the runoffs at the river mouths get distributed
uniformly all over the Bay even though salinity fluctuations over shelf areas caused by runoff are large compared with the open areas of the Bay. Also, for WM and SM
the net runoffs are only 0.1% and 0.7% of the oceanic transports
Table 3 Precipitation (cm/season and cm/yr) over the Bay.
Latitude Longitude Annual Mean(cm) Winter monsoon (Dec.+Jan.+Feb.) Summer monsoon (Jun.+Jul.+Aug.)
(°N) (°E) M Index (I) IxM Index (I) IxM
5–10 80–85 144 0.263 37.93 0.122 17.57
5–10 85–90 172 0.228 39.30 0.161 27.73
5–10 90–95 205 0.172 35.24 0.197 40.34
5–10 95–100 268 0.113 30.39 0.271 72.65
10–15 95–100 280 0.038 10.64 0.457 127.90
10–15 90–95 229 0.070 16.03 0.309 70.76
10–15 85–90 169 0.100 16.95 0.250 42.23
10–15 80–85 133 0.144 19.14 0.205 27.25
15–20 80–85 125 0.058 7.29 0.332 41.56
15–20 85–90 167 0.038 6.38 0.311 51.89
15–20 90–95 259 0.025 6.37 0.554 143.36
15–20 95–100 290 0.014 4.06 0.606 175.71
>20 90–95 290 0.019 5.45 0.627 181.95
>20 85–90 189 0.024 4.54 0.556 105.10
Table 4 Zonewise net water exchanges at the surface of Bay (WM=Dec.+Jan.+Feb., SM=June + July+Aug). (P=precipitation, R=runoff,
E=evaporation)
Latitude Longitude WM (cm) SM (cm) Net height (cm) Net exchange volume (×1010 m3)
(°N) (°E) P(+) R(+) E() P(+) (R+) (E) WM SM WM SM
5–10 80–85 37.9 3.7 38.9 17.6 34.0 40.0 +2.7 +11.6 +0.644 +2.768
5–10 85–90 39.3 3.7 39.4 27.7 34.0 41.1 +3.6 +20.6 +1.087 +6.221
5–10 90–95 35.2 3.7 37.6 40.3 34.0 43.9 +1.3 +30.4 +0.395 +9.233
5–10 95–100 30.4 3.7 39.9 72.6 34.0 34.8 5.8 +71.8 −1.380 +17.088
10–15 95–100 10.6 3.7 33.2 127.9 34.0 30.1 18.9 +131.8 −3.929 +27.400
10–15 90–95 16.0 3.7 29.3 70.8 34.0 33.3 −9.6 +71.5 −2.807 +20.910
10–15 85–90 17.0 3.7 35.6 42.2 34.0 40.2 −14.9 +36.0 −4.404 +10.639
10–15 80–85 19.1 3.7 37.3 27.2 34.0 36.8 −14.5 +24.4 −4.405 +7.413
15–20 80–85 7.3 3.7 36.5 41.6 34.0 34.9 −25.5 +40.7 −3.180 +5.076
15–20 85–90 6.4 3.7 31.6 51.9 34.0 30.4 −21.5 +55.5 −6.161 +15.904
15–20 90–95 6.4 3.7 26.5 143.4 34.0 34.1 −16.4 +143.3 −4.180 +36.521
15–20 95–100 4.1 3.7 22.6 175.7 34.0 27.4 −14.8 +182.3 −0.621 +7.644
>20 90–95 5.4 3.7 28.4 182.0 34.0 30.5 −19.3 +185.5 −1.089 +10.470
>20 85–90 4.5 3.7 34.2 105.1 34.0 26.8 −26.0 +112.3 −1.522 +6.574
Grand Mean (cm) −11.5 +63.7
Total −31.552 +183.861
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across the 6°N section between surface and 1250m minimizing the effect of runoff on total water balance (see pp. 56–62). Computations show that the net exchange
heights are −11.5cm and +63.7cm for WM and SM, respectively. Hence during WM and SM the Bay shows negative and positive water balances respectively.
Coastal oceanography
Sewell (1928) found average surface salinities varying from 33.27 to 33.81ppt during October 1921 to February 1922 in Nankauri harbour, Nicobar Islands. Sewell
(1929) reported that large quantities of fresh water enter the Bay through the Ganges and Irrawady rivers resulting in significant reduction in surface salinity in the
northern Bay. He observed that during the summer monsoon the greater part of the Bay surface is occupied by water of salinity less than 34.0ppt. But in Palk Bay
salinities between 34.5ppt and 35.0ppt were observed. Ganapathi & Murty (1954) observed surface salinities of about 29.0ppt in January, 34.0ppt in March off
Waltair and below 18.0ppt during the summer monsoon. Ganapathi et al. (1956) observed bottom salinities between 33.5ppt and 34.5ppt over the central shelf and
stated that the vertical salinity gradients reflected the runoff and circulation. Ganapathi & Rao (1959) reported high surface temperatures during JulyOctober, when
flow of fresh water into the Bay is maximal and low surface temperatures during DecemberFebruary when the discharge is minimal. These low and high temperatures
were attributed to surface cooling caused by low insolation.
La Fond (1954b) studied upwelling and sinking along the east coast of India from March to May and from September to November. Temperature oscillations
observed near the mouth of the River Krishna were attributed to internal waves (La Fond 1945a, La Fond & La Fond 1968). Oscillations with periods ranging from
12h to a few minutes were observed over the continental shelves of the Bay (La Fond 1962). La Fond & La Fond (1968) observed an offshore displacement of
surface water from the Burmese coast resulting in nearshore upwelling. Jayaraman (1954) observed that during MayOctober salinity in the Gulf of Mannar and Palk
Bay (near Mandapam) increased (maximum salinity of 37.0ppt) because of northward flowing currents from the Indian Ocean and Arabian Sea. During December
January salinities less than 23.9ppt were recorded. Ramasastry & Balaramamurty (1957) found that surface temperatures increased towards offshore during post
summer monsoon (OctoberNovember) except near the mouths of the Godavari and Krishna rivers. They also found that surface salinity decreased during October,
reaching a minimum by December. From December, salinity values increased to a maximum during March. Balaramamurty (1958) carried out observations of the time
and space variation of temperature and salinity structures of the western Bay of Bengal in general and the monthly mean conditions off Visakhapatnam in particular. At
greater depths the isotherms were closely packed due to intense upwelling from deeper layers bringing up colder waters. He further found that the month of maximum
intensity of upwelling varied between April and March off the coast of Visakhapatnam, although the period of upwelling generally extends until May. Varadachari
(1958) studied upwelling and sinking off Waltair.
Murty & Varma (1964) found that Bay of Bengal waters entering Palk Bay considerably influenced hydrographic conditions there. It was reported (Anonymous
1964, Radhakrishna et al. 1982) that in the northern Bay, with increased runoff during summer monsoon, salinity values off river mouths decreased to about 18 ppt, in
southern areas to about 34ppt through
Page 11
30ppt in the middle sector. La Violette (1967) found very low salinity values (<20ppt) near the mouth of the Irrawady river during the summer monsoon. He observed
the effect of runoff from the Ganges up to about 800km offshore and cooling of surface water along the east coast of India with monsoon winds and rains. Prevalence
of upwelling along the coasts of India and Burma was also noted during winter. The seasonal fluctuation of the surface temperature, salinity, dissolved oxygen and other
nutrient salts during 1964–66 indicated the occurrence of upwelling (AprilJune) and sinking (SeptemberOctober) off Waltair during certain months of the year
(Mojumder 1967). Anand et al. (1968) found that upwelling occurred off Madras and further north, off Visakhapatnam, during the summer monsoon. Rao &
Jayaraman (1968a) observed deepening of the halocline, pycnocline and thermocline and low saline water in the regions off the mouths of the Krishna and Godavari
rivers. Yesaki & Jantarapagdee (1981) found the water column over the continental shelf off the west coast of Thailand to be strongly stratified during the winter
monsoon and weakly stratified during the summer monsoon. Two regions of upwelling were observed (De Souza et al. 1981) near Madras and Visakhapatnam during
AugustSeptember. Antony et al. (1985) observed temperature oscillations at four different levels between 14 and 20m with an average periodicity of about 16h in the
vicinity of the River Krishna. Gopalakrishna & Sastry (1985) found indications of strong upwelling along the southern region off the east coast of India. Vertical
temperature and salinity structures were of estuarine nature in areas north of Visakhapatnam. Rao et al. (1986) found upwelling of cold water up to the surface from
February to May off Visakhapatnam at 60m depth.
In the following paragraphs property distributions, both along the vertical and horizontal, are described for summer and winter monsoons in the coastal areas (250–
300km) from the southwest Bay of Bengal to southeast Andaman Sea. Discussion sections are also added appropriately for each property.
Temperature fields
During the winter monsoon of 1983 surface temperature (Fig. 5) increased from 25.0°C at 18°N to 28.5°C towards the south of Madras and to 26°C at about 20°N.
At 50m the temperature varied from 26°C at 16°N to 28°C towards both the south and north. A few pockets of higher temperatures compared with the surface, were
observed at this depth. Balaramamurty (1958) and Rao and Sastry (1981) attributed this to surface cooling. At 100m during the winter monsoon a broad coldwater
cell (<17°C) appeared between 13°N and 16°N (Suryanarayana et al. 1993). At 200m also, below the thermocline, the lowest temperature (13°C) was recorded
around 16°N, increasing towards the south and north. During the winter monsoon the sea surface temperature (Fig. 10) in the southwestern region was high compared
with that in the other regions along the coast. The thermocline first dips to about 100km and then becomes shallower towards offshore. Off Visakhapatnam, the
thermocline is deeper and becomes uniformly shallower from 100 to 200m (Fig. 11). Further northward off Mahanadi (Fig. 12), the thermocline shows a similar
structure to that at Visakhapatnam. At the head of the Bay also, off the Burma and Bangladesh coasts, the surface temperature is 26°C (Fig. 13) as is that off Chilaka
Lake. Here, from the wide shelf, the isotherms dip towards deep regions. Off the Irrawady towards the west and south (Figs 14, 15) surface temperatures ranged
between 27.5°C and 28°C during March 1963. A sharp thermocline was noticed only in the westeast section (Fig. 14) and not in the northsouth section (Fig. 15)
where the maximum depth is only about 80m. During JanuaryFebruary of
Page 12
Figure 5 Distribustion of temperature during winter at various levels in coastal areas.
Figure 6 Distributions of salinity during winter at various levels in coastal areas.
Figure 7 Distributions of temperature during summer at various levels in coastal areas.

Page 13
1983, Janekarn & Hylleberg (1989) found low temperatures between 25.6°C and 27.2°C in the coastal waters off the west coast of Thailand. They found that bottom
temperatures varied greatly, in some places between 25°C and 26.7°C, and in other places between 27.5°C and 28.3°C. They also observed thermocline depths
varying between 40m and 45m in the coastal and offshore areas (from 9° 5′N to 6° 30′N) during JanuaryApril. These surface temperatures are about 2°C lower than
those observed off the Madras coast on the west side of the Bay. Similarly, these thermocline depths are also much lower compared with those observed between 18°
N and 21°N along the west coast of the Bay.
For the summer monsoon, data are available only along the east coast of India. Surface temperatures vary between 28°C and 30°C (Fig. 7). In the northern coastal
belt, temperature increases by 4°C (26°C to 30°C) from winter to summer, whereas off Madras temperatures are only around 28°C. At 50m and 200m surfaces,
pockets of high and low temperatures are seen, caused by fluctuations in the depth of the thermocline. Off Madras a shallow thermocline (about 50m) without any
marked upwelling is observed (Fig. 10). Off the central east coast of India (Fig. 11) the thermocline shows a wavy pattern. Similar wavy temperature structures were
observed by Babu et al. (1991) in the same area during summer. A warmwater cell (23°C) is seen off the central east coast of India at 100m (Fig. 20b see p. 29).
Over the northern coastal belt (Fig. 12) the thermoclines show little change from winter to summer but surface temperatures increase during summer. Sundstrom et al.
(1987) found surface temperatures of 29°C in the Andaman Sea near Phuket Island during AprilMay. This is almost the same temperature as that observed off the
southern coast of India (Fig. 7).
A sea surface temperature gradient from north (26°C) to south of Madras (27°C) during winter is caused by the cold, dry northeasterly winds aiding latent heat flux
(evaporative cooling) and sensible heat flux (convection etc.) from the sea surface in the northern Bay of Bengal. Surface temperatures in the southern Bay (5–10°N
and 85–95°E) remain uniformly high between 28.0°C and 28.5°C during winter (see pp. 24–35). But in the Andaman Sea from about 96°E to the shelf areas (about
100m depth) the surface temperatures are found to be 28°C. Shetye (1993) noted that the GangesBrahmaputra discharge decreased the sea surface temperatures in
the northern Bay during the winter monsoon. But from Figures 5, 6, 7, 8, 18, 20 (see p. 29), 27 (see p. 37) and 29 (see p. 39), it seems that river runoff has no
significant effect on the surface temperatures compared with winter cooling. The lowtemperature (<25°C) cell observed at 50m (Fig. 7) during summer represents a
cold core eddy. A similar eddy was reported by Babu et al. (1991). They found that the eddy has a temperature drop of 4–5°C at its centre and extends between
50db and 300db surfaces with a diameter of about 200km. Eddies are generally generated by baroclinic instability at the interface of two opposing currents present
along the shelf edge. In the northern Andaman Sea at Station 26 (Fig. 14) a temperature inversion is observed. Similar inversions are also observed along the east coast
of India during winter (Fig. 5). These inversions are considered to be of high significance in the Bay. The thickness of the mixed layer extends up to 60m outside the
shelf area in the northern Andaman Sea.
Salinity fields
During the winter monsoon, surface salinities range between 27.0ppt and 33.0ppt (Fig. 6). The 33.0ppt isohaline runs almost parallel to the coast. Lowest salinity
(27.0ppt) is observed in the region around 20°N and is a result of river runoff from the GangesBrahmaputra system. At 50m the 33.0ppt isohaline breaks at different
regions and high saline waters
Page 14
Figure 8 Distributions of salinity during summer at various levels in coastal areas.
Figure 9 Map showing positions of various stations used for vertical hydrographic sections.
Page 15
Figure 10 Vertical sections of A, salinity, B, temperature and C, density along SM1 (top) and WM1 near Madras for summer and
winter, respectively.
Figure 11 Vertical sections of A, salinity, B, temperature and C, density along SM2 (top) and WM2 near Visakhapatnam for summer
and winter, respectively.
reach nearshore areas. At 100m, over most of the shelf, salinity varies between 34.6ppt and 34.9ppt except near the river mouths where the effect of freshwater
discharge is still noticed (Suryanarayana 1988, Suryanarayana et al. 1992). At 200m salinity is more uniform with insignificant mixing with top diluted water (<34.5ppt).
At 500m it varies from 35.05 to 35.10ppt (Suryanarayana 1988, Suryanarayana et al. 1991). During the winter monsoon along 14°N (down—A, Fig. 10) surface
salinities vary from 32.0 to 33.4ppt from about 100 to 300km offshore compared with 27.0 to 32.5ppt along 20°N from 50 to 300km offshore (down—A, Fig. 12).
The diluted water (<34.5ppt) is observed only up to 40m depth (Fig. 10). Along this section the salinity contours tilt upwards offshore (at about 200km) to the surface.
Off the central coast at about 17°N surface salinities (Fig. 11) vary between 27.0ppt near the shore and 33.5ppt (about 200km). Diluted water extends up to 200km
and vertically
Page 16
occupies about 60m of the water column. During the same season in the northern region (20°N) (down—A, Fig. 12), surface salinities vary from 27.0ppt near the
shore to 32.5ppt towards offshore (300km). The diluted water (<34.5ppt) extends up to over 300km from the shore and up to 50m of depth. Surface salinities are >
30.0ppt at the surface off the river Ganges for about 200km (Fig. 13). Here the surface diluted water extends up to about 75m depth. In the Andaman Sea surface
salinity increases from 32.0ppt in the north (Fig. 14) to 33.0ppt in the south (Fig. 15). Janekarn & Hylleberg (1989) also found surface salinity variations between 32.
1ppt and 33.6ppt in the coastal waters off the west coast of Thailand during JanuaryFebruary of 1983. The diluted water (<34.5ppt) occupied the full shelf areas over
the northern and eastern regions. This water column extends up to about 75m over the full northeastern Bay (Fig. 13 & 14).
During the summer monsoon, surface salinities range between 22.0ppt and 34.0ppt over the coastal area (Fig. 8). The 33.0ppt isohaline forms a boundary across
the central east coast (at about 16°N) between the southern high salinity belt and the northern low salinity belt. The low salinity front in the north is formed as a result of
heavy runoff from the MahanadiGangesBrahmaputra systems (see pp. 7–8). Off Madras (top—A, Fig. 10) the diluted water extends up to about 50m with more or
less uniform thickness, whereas in winter the pattern shows wavy structures. Off Visakhapatnam (around 17°N) the diluted water extends up to about 70 to 90m with a
wavy pattern across the shelf (top—A, Fig. 11). In the northernmost section (top—A, Fig. 12) the diluted water extends up to >110m with almost uniform thickness
(surface salinities shown in Figure 12 are higher compared with those in Figure 8 due to presentation limitations). A similar increase of thickness of the layers of diluted
water was noted also by Shetye et al. (1991). No results are available for the coastal areas of the northeastern Bay of Bengal and Andaman Sea for summer. Although
the lowest salinity in Figure 8 is 22.0 ppt, further northwards salinity decreases to 21.0ppt (Fig. 21a, see p. 30). Surface salinities (low) in this region can be expected
to change from year to year and within seasons. Gopalakrishna & Sastry (1985) reported a low salinity of 20.0ppt. Another low value of 16.0ppt was reported by
Murty et al. (1990).
Figure 12 Vertical sections of A, salinity, B, temperature and C, density along SM3 (top) and WM3 near Mahanadi for summer and
winter, respectively.
Page 17
Figure 13 Vertical sections of temperature and salinity along section WM4 off Ganges for winter (reproduced from Maslennikov 1973).
Figure 14 Vertical sections of salinity, temperature and density along WM5 off Irrawady for winter (after Ramesh Babu & Sastry
1976).
The range variation of surface salinities during summer is twice (12ppt, from 22.0 to 34.0ppt) that of winter (6ppt, from 27.0 to 33.0ppt) (see p. 7 for runoff estimates
in summer and winter). The surface diluted water (< 34.5ppt) shows wavy distribution patterns (in thickness) across the shelf irrespective of the season. Compared
with the large range of fluctuations in the coastal areas, the deep Bay experiences only low ranges of fluctuations; 29.0
Page 18
to 34.5ppt during summer and 31.4 to 34.0ppt during winter (see Figs 27 p. 37 and 29 p. 39). The lowest deep Bay salinities are observed along 15°N and are
caused by the dilution effect from river runoffs. The surface salinity distributions are found to follow the seasonally changing current patterns. The low salinity tongue
extending down to Visakhapatnam from the north is caused by the southward flowing coastal part of the gyre in the northwestern Bay (see Fig. 42 p. 58). The high
salinities in the southern region are caused by the northward extension of the Indian Monsoon Current. During winter, relatively low salinity waters (see Fig. 29) flow
into the Bay through the North Equatorial Current; this mixes with the diluted surface water and recirculates it through the two clockwise gyres (see Fig. 42) further
reducing salinities in the interior Bay. This seems to be the reason for the continuous low salinity coastal belt. In general salinity and temperature distributions over the
coastal belt are controlled by the circulation, which in turn is caused mainly by wind and freshwater discharges through the MahanadiGangesBrahmaputra system at
the head of the Bay and the KrishnaGodavari system at the central east coast. The horizontal distributions show the existence of frontal areas due to river runoff,
coastal currents, upwelling etc. The persistence and proportions of such fronts are complicated (Csanady 1978).
Figure 15 Vertical sections of temperature, salinity and density along WM6 for winter (after Ramesh Babu & Sastry 1976).
Page 19
Density fields
During the winter monsoon, sea surface density varies between 17.0 and 21.0 and the isopycnals run nearly parallel to the coast (Suryanarayana 1988). Sigmat at
100m ranges between 22.0 and 25.0 across the shelf. Pockets of high (low) sigmat coincide with saline (diluted) water cells at surface. Off Madras (down—C, Fig.
10) the 21.0 sigmat isopycnal encloses a low density water body. Off Visakhatpatnam (down—C, Fig. 11) and Chilaka Lake (down—C, Fig. 12), very low density
water bodies are observed near to the coast. Along the coast from Madras to Chilaka Lake nearshore subsurface isopycnals surface about 100km offshore. In the
northern Andaman Sea (Fig. 14) the shelf water seems to be well mixed. Along the east coast of the Andaman Sea, the water column is mildly stratified with a weak
discontinuity at about 40m (Fig. 15).
During the summer monsoon, sigmat ranges between 12.0 (off Mahanadi) and 21.5 (off Madras) and the nearly parallel nature of the isopycnals, observed during
winter, is absent (Suryanarayana 1988). The effect of freshwater dilution is very different, compared with that noted in winter, over the northern region. Over 100m
surface sigmat ranges between 23.0 and 24.5 from north to south (Suryanarayana 1988). During summer the isopycnals (Figs 10, 11, 12) do not slope up towards
offshore as observed during the winter. The 21.5 sigmat isopycnal (across the shelf) shows a gentle shallowing from about 70m in the north (Fig. 12) to about 30m in
the south (Fig. 10).
During both winter and summer monsoons (Fig. 10), high densities (>20.5) prevail over the southern region at the surface compared with northern parts of the
coastal belt. Freshwater outflows are seen in the form of closed isohalines or low salinity in the vicinity of the river mouths (Figs 6, 8). The isohaline labelled 34.5ppt
separates the upper diluted water from the high salinity deep water. The low range variations of salinity in the southern Bay (near 11°N) show that the influence of fresh
water is largely limited to northwestern and northeastern regions. A stretch of coastal sea 150 to 200km wide and about 50m deep northwards from about 15°N can
be treated as the zone where fresh water plays a very important role in the dynamics. The KrishnaGodavari system brings in 1.1×109m3 per month during the winter
and 29.8×109m3 per month during summer (Table 1). Similarly, the MahandiGangesBrahmahputra system brings in 18.7×109m3 per month and 156.6×109m3 per
month during winter and summer, respectively.
The northern Bay of Bengal and Andaman Sea are also very strongly affected by continental runoff from the IrrawadySalween system. This systems outflows
28.3×109m3 per month and 307.7×109m3 per month during winter and summer, respectively (Table 1, see p. 7). For this region, only very limited information is
available on density (Figs 14, 15), but from Figure 29 (see p. 39), based on data from deep sea cruises, it is clearly seen that the effect of runoff (marked by the
33.0ppt isohaline) extends up to 90°E and 9°N. Maslennikov (1973) noted that the vertical distributions of temperature and salinity in the northeastern part of the Bay
of Bengal are similar to their distributions in the Andaman Sea down to 700–800m.
Upwelling
One of the unique features of the Bay of Bengal and Andaman Sea is upwelling and sinking along the coasts. Murty & Varadachari (1968) reported strong upwelling
off the Waltair coast (with waters from depths greater than 100m reaching the surface) and weak upwelling
Page 20
Figure 16 Distribution of upwelling indices off various stations along the east coast of India. Units are in m3sec−1 100m−1 coastline. See
Figure 9 for Stations marked 1 to 8.
(waters from 30m depth reaching the surface) off Madras during 1964. The difference in intensity of upwelling during this period was attributed by them to the relatively
strong winds along the coast of Waltair (see Fig. 4). The currents along the coast of Madras are reported to be variable during June and sometimes they even flow
towards the south (see Fig. 42, p. 58).
Page 21
Bakun (1973) defined an “upwelling index” (UI) as the magnitude of the offshore component of the Ekman transport (Mx=τy/f) where τy is the wind stress and f the
Coriolis parameter. The UI is considered to be an indicator of the amount of water that upwells through the bottom of the Ekman layer to replace water driven offshore
by the longshore wind stress. Figure 16 (Suryanarayana 1988) shows this UI at eight coastal stations (see Fig. 9 for the locations of stations numbered 1 to 8) along the
east coast of India. From this figure it is clear that there is strong upwelling during the summer monsoon and downwelling during winter months. Based on the
distribution of UI, the entire east coast of India can be divided into three zones, northern (Stations 1 and 2), central (Stations 3, 4 and 5) and southern (Stations 6, 7
and 8). Such a zonation is supported by the salinity fronts (Figs 6, 8) that are caused by the combined effects of MahanadiGanges river flows. Again, Stations 3, 4 and
5 fall in the area of the coastal part of the northwestern gyre (see Fig. 42 p. 58). In the northern zone UI values are negative. In the central zone UI values are low
compared with those of the southern regions. At Station 2, negative values are high in comparison with those at 3 and 4. The intensity of positive and negative indices
are found to be equally distributed, however, at Stations 3 and 4. At Station 5 high positive values were noted during 1983 and 1984, however, showing high wind
induced upwelling. It seems that during these two periods the upwelling belt extended abnormally northwards because of wind or currentinduced factors. In the
southern zone high values of UI were noted compared with other northerly zones with large fluctuations. Stations 6, 7 and 8 are in an area of intense offshore surface
currents (Fig. 42) and hence favourable for upwelling. Also, from Figure 16 it is clear that upwelling shows variations from year to year especially at Stations 5, 7 and
8. Figures 10, 11 and 12 also reveal variations from south to north and between seasons. The progressive decrease of UI values from south to north is caused by the
changing coast alignment and changes in wind direction and speed (see Figs 3, 4 p. 5–6). In reality the windinduced upwelling would be influenced by currents flowing
in from remote regions like the North Equatorial Current (Fig. 42, p. 58) during winter.
Hydrographic data collected (Shetye et al. 1991) during the summer monsoon of 1989 along the east coast of India showed (a) an upwelling band (about 40km
wide) along most of the coastline, (b) cellular structures in dynamic topography offshore of the band and (c) a southward moving freshwater plume over the
northwestern Bay of Bengal. The upwelling band is apparently driven by the local winds alone. The geostrophic velocity in the band was about 50cms−1 in the
poleward direction. Below the upwelling band, downwelling was often observed suggesting the presence of an undercurrent. Off Waltair this upwelling band was a few
miles wide in March, and by May it covered half the width of the shelf. In the southern part of the Andaman Sea the vertical distributions of hydrological parameters at
the centre of the anticyclonic gyre exhibit a band of isohalines indicating sinking of the surface water (Maslennikov 1973). Strong upwelling is caused by a
compensatory undertow of the anticyclonic gyral periphery. Similar upwelling occurs in the northeastern part of the Bay of Bengal, where an anticyclonic movement is
noticed. Figures 10, 12 and 13 show the movement of deep, nutrientrich, cold and saline waters on to the shelf during winter, off Madras, Mahanadi and off the
Burma coast, respectively. The distribution of temperature, salinity and values of vertical stability in the northern part of the Andaman Sea also indicate upwelling over
shelf waters. Yesaki & Jantarapagdee (1981) noted that upwelling over the continental shelf off the west coast of Thailand is primarily windinduced.
Page 22
Turbidity and light

La Fond & Sastry (1957) used a hydrophotometer to study the transparency of the coastal waters along the east coast of India. They found that in the summer season,
turbid water tends to remain nearshore and flows down the coast. As the rainy season recedes the water becomes clearer in nearshore areas, remaining, however,
more turbid than that over the shelf. During OctoberNovember the transparency or percentage of light transmitted with respect to air in the nearshore was about 60 to
90% in the surface layer and less than 20% near the bottom. The transparency at the surface in January and February was 80 to 100%. In March the surface remained
clear but the upwelled subsurface layers near to the shore became turbid. In April the upwelling reached the surface and the plankton bloom was heaviest affecting the
transparency of the water column. From the observed pattern of turbidity in the sea La Fond & Sastry (1957) tried to explain the water structure and circulation off the
east coast of India. Satyanarayana Rao (1957) found that during November and December 1952 the waters all along the east coast of India showed uniform
transparency. Sundara Raman & Sreerama Murty (1968) found that during the month of March the transparency was 40 to 60% up to 8km off the Karaikal coast. It
increased to 100% further offshore. The transparency off Madras was uniform and low and towards offshore it increased to 50%.
The turbidity of the waters off the east coast of India vary greatly from season to season depending on runoff and associated suspensions. Suspended sediments
enter the Bay of Bengal through the many rivers, mainly during the summer monsoon. The Bay receives about 16×108 tonnes of silt yearly (Suryanarayana 1988) along
with abundant runoff (see Table 1, p. 7). The suspended materials settle down or get transported away from the river mouths depending upon currents. As there are
large seasonal variations in currents and runoff, transparency also shows great changes.
Deep bay oceanography

Sewell (1929) documented the seasonal surface salinity distribution and surface currents in the Bay of Bengal and Andaman Sea. His analysis showed large meridional
gradients of salinity during SeptemberNovember and low meridional gradients during MarchApril, coinciding respectively with the periods of highest and lowest
freshwater influx from the Ganges and Irrawady rivers. He pointed out that the pattern of surface currents and surface salinity distributions are related. Sewell (1932),
for the first time, presented the seasonal variation of temperature in the depth interval 300–1000 fathoms (549–1829m) in the Bay of Bengal and Andaman Sea and
found that at all depths the rise and fall of temperature follow an almost identical curve, but the changes in the Andaman Sea appeared to lag behind those in the Bay of
Bengal by an interval of about a month. He noticed comparatively higher changes of temperature at 100 fathoms (183m) during OctoberNovember in the Bay of
Bengal. His analysis further showed that deep (>900m) waters of the Andaman Seas were warmer than those of the Bay of Bengal and the difference in temperature
was as high as 1.7°C at 1000 fathoms (1829m). Wyrtki (1961) studied the physical oceanography of the southeast Asian waters and identified a subsurface (75–
100m) high salinity maximum just below the surface mixed layer in the equatorial region of the eastern Indian Ocean that he attributed to the penetration of Arabian Sea
high salinity water. While examining the structure of the water masses of the Indian Ocean, Rochford (1964) identified a high salin
Page 23
ity water mass in the Bay of Bengal at subsurface depths (75–100m) and a high salinity layer in the range of 200–400m. Ivanov (1964) studied distributions of
temperature and salinity in the upper levels of the Bay of Bengal and the Andaman Sea. Gallagher (1966) discussed the circulation and structure of water masses of the
Bay of Bengal. Utilizing the hydrographic data available prior to 1967, La Violette (1967) presented the seasonal distributions of temperature, salinity and density in the
Bay of Bengal and Andaman Sea. Varadachari et al. (1967) observed variations of thermocline depths from 20 to 8m in the western Bay of Bengal. Anand et al.
(1968) presented the temperature and oxygen distributions on various horizontal surfaces in the upper 300m of the west central Bay of Bengal during the summer
monsoon (JuneJuly) of 1964. The pattern of isopleths of oxygen was found to be similar to that of temperature during this season. The concentration of oxygen in the
oxygen minimum layer and the thickness of the oxygenrich surface layer were found to be least (minimal) in the Bay of Bengal compared with those in the Arabian Sea
and equatorial region. Rao & Jayaraman (1968a) presented temperature, salinity and density distributions along various zonal sections and discussed the water mass
structure and circulation in the central and southern Bay of Bengal during FebruaryMarch 1963. Wyrtki (1971) prepared a comprehensive and voluminous
oceanographic atlas of the Indian Ocean based on the IIOE data.
Colborn (1975) compiled the temperature data of the Indian Ocean and presented the thermal structure in a detailed way in various areas including the Bay. Ramesh
Babu & Sastry (1976) presented the distributions of temperature, salinity, density and dissolved oxygen during March 1963 in the Andaman Sea and discussed the
field of motion in relation to the influx of fresh water. They reported high (4.6mll−1) dissolved oxygen at the surface and large variation of dissolved oxygen at 50m and
100m which they attributed to the convergences and divergences in the field of motion. Rao (1977) investigated the physical processes that control the potential
productivity of the Bay of Bengal and the Arabian Sea. His study revealed that the relatively weak upwelling and high degree of stratification caused by the river
outflows in the Bay of Bengal affected the transfer of nutrients across the thermocline from the subsurface depths to the sea surface. Rao & Sastry (1981) showed that
the integral mean concentrations of nutrients in the surface mixed layer was in consonance with the nearsurface circulation. They noted that at the centre of the cyclonic
cells the integral mean concentrations of inorganic phosphates were high, whereas at the centre of the anticyclonic cells the values were low. Rao et al. (1981) studied
the shortterm variability of mixed layer depth in the northern Bay of Bengal. Varkey (1986) studied evaporation, precipitation, runoff, water transports and mixing of
different water masses for the whole Bay for the winter monsoon.
Murty (1990) investigated the large scale circulation in the Bay of Bengal during the summer monsoon (JulySeptember) of 1984. His analysis revealed that the
thermohaline structure and the circulation in the Bay of Bengal are very much influenced by freshwater influx, wind forcing and the Indian Monsoon Current that enters
the Bay from the southwest. Recently many studies on water characteristics and circulation in the Bay of Bengal have been carried out for the summer monsoon (Murty
& Rao 1992, Murty et al. 1992, 1993, 1994), early winter monsoon (Suryanarayana et al. 1993) and presummer monsoon based on data collected on board
research vessels RV GAVESHANI and ORV SAGAR KANYA.
In the foregoing paragraphs, distributions of temperature, salinity and sigmat (or thermosteric anomaly in a few cases) at the sea surface (100m, 200m, 500m
1000m, 2000m and 3000m) for four seasons are presented and interpreted (see p. 3 for various groupings of seasons) based on data and results published earlier. The
figures were carefully selected to
Page 24
represent prevailing water properties. Some typical vertical sections are also used for the discussion.
Temperature fields
Surface
Figures 17a, 18a, 19a and 20a represent the distributions of sea surface temperature (SST) in the Bay of Bengal during the four seasons. During the presummer
monsoon season (Fig. 17a) SST increases from 27.5°C in the north to 30.5°C in the central Bay. The SST distribution indicates the warming of the surface waters
between winter and summer. The lower SSTs in the northwestern Bay show the aftereffects of strong surface cooling during winter. The higher SSTs in the central Bay
represent the seasonal peak during presummer monsoon. Climatological data (Verploegh 1960, Colborn 1975, Hastenrath & Lamb 1979) and data from various
cruises (Somayajulu et al. 1987) during the presummer monsoon months show higher SSTs exceeding 30°C up to 30.5°C in the central Bay. The AVHRR SSTmaps
of the Bay also show SSTs exceeding 30°C up to 31°C in the central Bay for AprilMay (Viswambharan & Mathew 1992, Murty et al. 1994). In the northeastern
Bay, SSTs exceeding 28°C were reported during AprilMay 1963 (La Fond & La Fond 1968) with relatively colder (<27°C) waters close to the Burma coast under
the influence of coastal upwelling from a depth of about 100m. They also noted the occurrence of a clockwise eddy of about 240km in diameters centred around 18°N
and 88°E that affected the thermal structure in the northwestern Bay. These authors further reported high SSTs varying between 29°C and 30°C in the western Bay off
the central east coast of India during AprilMay 1963, and high SSTs exceeding 30°C outside the upwelling zones off the coasts of Visakhapatnam and Madras.
During the summer monsoon (Fig. 18a), warm (>29.5°C) waters are noticed in the northwestern Bay. From this region, SST decreases gradually to 28°C towards
the southwest along the Indian coast and southeast in the offshore region. The surface waters of the central Bay have a temperature around 28.5°C over a large area
although relatively higher (29°C) and lower (28°C) SSTs are also observed at some stations. In general, it can be seen that the surface waters south of 14°N are cooler
(<29°C) compared with those north of 14°N. The southern cold surface waters indicate the influence of summer monsoon in lowering the SSTs by the drastic decrease
of insolation, increase of latent heat flux across the air/sea interface and enhanced turbulent mixing resulting from strong winds. The northern warm surface waters are
clearly due to high insolation in the north Bay during July. Earlier Anand et al. (1968) reported SSTs varying between 28.5°C and 29.3°C in the central Bay with
warmer waters in the west during June 1964 and lower SSTs (<28°C) close to the coast off Visakhapatnam during July 1964.
During the postsummer monsoon (Fig. 19a) distribution of SST is characterized by a cell of warm water with a core value of 29.5°C centred at 8°N: 85°E. Lower
(<28.5°C) SSTs are seen in the eastern Bay of Bengal. Somayajulu et al. (1987) noticed a temperature variation of 0.5°C from 28.3°C to 28.8°C along two transects
(orientated northwestsoutheast) in the central Bay of Bengal during the postsummer monsoon period of 1983. Murty et al. (1992b) observed warm (28.5–29.5°C)
SSTs in the northwestern Bay of Bengal during the retreat of the summer monsoon (September) of 1983.
During the winter monsoon (Fig. 20a), the lowest SST is 25.5°C observed in the northern Bay and SSTs around 26°C are seen off the central east coast of India. In
the southern Bay and Andaman Sea, SST fluctuates between 28°C and 28.5°C. In the southern and central Bay
Page 25
Figure 17 Horizontal sections of temperature for presummer monsoon at 0m, 100m, 200m and 500m for Bay of Bengal.
variations of SST between 26°C and 28.6°C during FebruaryMarch 1963 were reported by Rao & Jayaraman (1968a) and the higher SSTs (>27.5°C) were
attributed to the diurnal sea surface temperature maximum. The temperature difference from the head of the Bay to the southern Bay is 3.0°C, which is similar to that
documented earlier (Balaramamurty 1958, Wyrtki 1971). The general southward increase is partly accounted for by latitudinal variation
Page 26
Figure 18 Horizontal sections of temperature for summer monsoon at 0m, 100m, 200m and 500m for Bay of Bengal.
of insolation. The high SST values in the Andaman Sea also seem to be the result of high insolation. The SSTs in the northern Bay indicate the effect of whiter cooling
with more cooling off the Burma coast under the influence of cold, dry northeasterly winds blowing from the land into the Bay of Bengal. La Violette (1967) also
noticed a sharp decrease in SST in the extreme northern Bay of Bengal from NovemberDecember to JanuaryFebruary and this was attributed to a “decrease in net
surface insolation although increased evaporation due to
Page 27
the dry winter monsoon winds could also aid surface cooling”. The effect of cold river discharge on the surface temperatures can be noticed off Visakhapatnam and the
central east coast of India. In the northwestern Bay, cold (<26.0°C) surface waters and a temperature inversion of 1°C at 50m at the distance of 360km from the
Gopalapur coast were reported during January 1965 (Sankaranarayana & Reddy 1968). Rao & Sastry (1981) reported temperature inversions of 1.5°C at depths of
50m in the northern Bay during January 1963. Suryanarayana et al. (1993) also reported low (<26.5°C) temperatures at near surface depths and a temperature
inversion of 2°C at 50–75m in the northwestern Bay during winter 1983. The sea surface thermal field, derived from NOAA/AVHRR satellite data during December
February 1992, also shows low SSTs (<25.5°C) at the head of the Bay of Bengal (Murty et al. 1994). Colborn (1975) suggested downward transfer of heat as one
plausible mechanism for inversions. Another very possible mechanism seems to be the presence of major circulation gyres and small migrating eddies (see pp. 56–62).
From the above descriptions it can be seen that whereas during winter SST increases (3°C) from the head of the Bay towards the south, during summer it decreases
(1.5°C). Furthermore the effect of cold river runoff on lowering SST seems to be very small compared with insolation and evaporation and limited to areas just off river
mouths.
100 metre surface

Figures 17b, 18b, 19b and 20b show the distributions of temperature at 100m for various seasons. During the presummer monsoon, temperature variations show a
range of 11°C varying between 17°C and 28°C (Fig. 17b) compared with 3°C at 50m. A large warmwater cell is seen centred at 14°N and 85°E with a core
temperature of 28°C. Two coldwater cells with closed isotherms are noticed towards the west (18°C) and north (17°C) of the warmwater cell. Strong horizontal
temperature gradients prevail all over the area due to variations in the mixed layer. This temperature distribution indicates a strong poleward flow in the western Bay
and eastward flow centred along 17°N.
During the summer monsoon, temperatures range between 20°C and 26°C (Fig. 18b). A broad cold (20°C) water band, oriented in a southwestnortheast
direction, in the central Bay, characterises the temperature distribution at this depth. Such a coldwater band of about 24°C is seen also at about 50m. Towards the
east of this band, temperature increases up to 26°C and to 24°C in the northwestern Bay. Anand et al. (1968) also reported alternate warm (25–27°C) and cold
(17–18°C) water cells at this depth in the western Bay during the summer monsoon of 1964.
During the postsummer monsoon, temperatures vary between 19°C and 25°C (Fig. 19b). Many warm and coldwater cells are present at this depth. A warm
water pocket (25°C) is located at 12°N and 83°E. A bathy thermograph and hydrographic temperature profile at this location showed a temperature inversion centred
at 100m. Murty et al. (1992b) noticed a temperature variation of 3°C from 17°C to 20°C in the northwestern Bay during the retreat of the summer monsoon
(September) of 1983.
During the winter monsoon, temperatures vary between 18°C and 27°C (Fig. 20b) with strong horizontal temperature gradients in the northern, northwestern and
southeastern regions. The thermal structure is complex with the presence of warm and coldwater cells. Penetration of warm (27°C) waters from the southeastern Bay
towards the central Bay is evident. Cells of cold water (<22°C) were noticed off the central east coast of India and the southeastern Sri Lankan coast.
La Violette (1967) from his trimonthly maps of temperature at 100m showed warm
Page 28
Figure 19 Horizontal sections of temperature for postsummer monsoon at 0m, 100m, 200m and 500m for Bay of Bengal (after Rao
1977).
(24.4°C) waters in the central Bay during JanuaryMarch that shifted northwards during AprilJune, relatively warmer (22°C) waters in the central Bay during July
September, and the occurrence of cold (8.9°C) waters in the central Bay during October December. The wide range of temperature fluctuations at this depth, within
seasons, and between seasons is due to variations in mixed layer depths. Rao (1977) found the surfacelayer thickness to vary between 30m and 100m during winter in
the Bay.
200 metre surface

During the presummer monsoon (Fig. 17c), temperatures vary between 12°C and 18°C. The large warmwater cell seen at 100m in the northern central Bay, together
with the coldwater cells in the northern Bay and western Bay (Fig. 17b), persist at this depth with core temperatures of 18°C, 12°C and 13°C, respectively. Relatively
large lateral temperature gradients prevail at the northern boundary between the warm and coldwater cells.
During the summer monsoon (Fig. 18c), temperatures vary over a narrow range between 13°C and 16°C. The well developed coldwater band at 100m is less
conspicuous at this depth, with weak lateral gradients from the central Bay to the western Bay. However, a large warmwater (>16°C) cell occupies the central Bay. A
small coldwater (<13°C) cell off Visakhapatnam coast separates the northern and central warmwater (15°C) cells. The temperature distribution indicates a cellular
mode of circulation in the central Bay, western Bay and along the east coast of India. During the postsummer monsoon (Fig. 19c), the temperature variation is 3°C,
from 13 to 16°C, with large warmwater cells in the central and southern parts.
During the winter monsoon (Fig. 20c), temperatures at 200m vary between 13°C and 16°C with warm water (16°C) cells in the central and northern regions. These
warmwater
Page 29
Figure 20 Horizontal sections of temperature for winter at 0m, 100m, 200m and 500 m for Bay of Bengal (after Rao 1977).
cells together form a warmwater ridge orientated approximately in a northeasterly direction. This indicates a poleward flow in the western Bay and an equatorward
flow in the eastern Bay. Temperature distributions at 200m are similar for the presummer, summer and winter monsoon seasons, with cellular patterns. However, the
coldwater valley (during summer) and the warmwater ridge (during winter), both oriented northeasterly, are characteristic of the temperature distribution at this depth.
Page 30
Figure 21 Horizontal sections of temperature for the four seasons at 1000m (winter plot after Rao 1977).
500 metre surface

At this depth, variations of temperature are between 9.0°C and 10.4°C during all seasons (Figs 17d, 18d, 19d, and 20d). Cellular patterns with small gradients are still
observed. Figures 23 and 24 show interrelated aspects at upper and lower surfaces.
Page 31
Figure 22 Horizontel sections of temperature, salinity and density at 2000m and 3000m.
Page 32
1000 metre, 2000 metre and 3000 metre surfaces

Figure 21 shows the seasonal distributions of temperatures at 1000m. The temperature variations are between 6.2°C and 7.0°C. The temperatures are higher by about
0.4°C in the northwestern Bay during the summer monsoon than those of the presummer monsoon. The waters in the central Bay are relatively warmer (6.8°C) during
the winter monsoon (Fig. 21d) compared with the other seasons. Cells of warm (7.0°C) water appear to persist along 12°N between 83°E and 85°E from summer
monsoon to the postsummer monsoon seasons. Figures 22a and 22d represent the distributions of temperature at 2000m and 3000m, respectively. Since the variation
in temperature is small, the isotherms are contoured at intervals of 0.1°C. At 2000m, the temperature varies between 2.6°C and 2.9°C and at 3000m the variation is
between 1.7°C and 1.8°C. In the following three paragraphs three vertical sections of temperature in the Bay of Bengal and Andaman Sea are presented for a
perspective study in relation to the horizontal charts.
Figures 23a and 24a represent typical meridional and zonal distributions of temperature along 91°E and 15°N, respectively, during the winter monsoon. Along 91°E,
a northward decrease of SST is evident owing to prevailing winter conditions in the northern Bay (see also Fig. 20a). The steep slope in the upward rise of the isotherm
(28°C) at about 7°N corresponds to a frontal area attributable to the westward flowing Northern Equatorial Current. The thermocline is strong south of 5°N. The
cellular patterns observed at 100m and 200m surfaces (Fig. 20) are the result of oscillations in the thermocline layer. Along the 15°N section, warm surface (>27°C)
waters are noticed east of 90°E (Fig. 24a). The surface layer ranges between 50m and 70m. It is shallow at 81°E (close to the east coast of India) and deep at 84°E.
Below the mixed layer, the isotherms exhibit a wavy nature with ridge and trough structures. In the western part, the isotherms indicate a northward flow between 82°E
and 84°E. Below 150m, under the northward flow, spreading of the isotherms can be seen, suggesting that there is an opposite flow in the depth range 150–300m to
the surface flow. At
Figure 23 Vertical sections of temperature along WM7 (a) and SM4 (b) from head of Bay to equator during winter and summer.
Page 33
greater depths, the isotherms are horizontal with much less eastwest variation of temperature (see also Figs 21, 22).
Figures 23b and 24b represent two typical meridional and zonal distributions of temperature in the central Bay along 89°E and 13°N, respectively, during the
summer monsoon. Along 89°E, the surface layer is more homogeneous in temperature and mixed layer thickness compared with winter (Fig. 23a), with its thickness
decreasing northward between 8°N and 15°N. The thermocline deepens gradually northward from 8 to 12°N and shoals gradually between 12°N and 18°N. The
deepening of the thermocline can be related to the convergence of water at subsurface depths which gets its support from the presence of a large warmwater cell at
200m in the central Bay (Fig. 18c). The shoaling of the thermocline and the lower SSTs between 15°N and 18°N are due to the subsurface divergence under the
influence of a cyclonic flow (Murty et al. 1992b) at the sea surface (see Fig. 42 p. 58). Between 300m and 450m the northsouth temperature variation is small.
Between 450m and 1000m the water column again becomes active. Along 13°N, the surface layer temperature is above 28°C between 81°E and 85°E (Fig. 24b).
The mixed layer is thinner than that of the winter layer across the Bay. The upward slope of isotherms below the surface layer between 85°E and 81°E indicates
upwelling on the Madras coast with a geostrophic northward flow. Between 86°E and 90°E, a well defined thermocline spreading between 50m and 300m is seen.
Because of doming of the isotherms, the surface layer becomes thin (40m). This spreading of the thermocline suggests a subsurface divergence zone centred at about
100m between 87°E and 88°33′E. Some recent data from the central Bay of Bengal during summer also show spreading of the thermocline in the central parts around
88°E. This coincides with the penetration of Arabian Sea high salinity water in the depth interval 75–100m (Murty et al. 1992a, Gopalakrishna et al. 1996). While the
doming of the isotherms above 100m is due to surface divergence caused by the prevailing positive wind stress curl field over this region (Babu 1987, Murty et al.
1992b) the down sloping of the isotherms below 100m may be due to the convergence of waters (see also Figs 18c, 23b) complementary to the surface divergence.
Below 500m, the horizontal variations of temperature are considerably less as seen earlier.
Figure 25 represents the meridional distribution of temperature along 95 °E (section WM9 in Fig. 9, see p. 14) passing through the central Andaman Sea during
winter. The surface waters are warm (>28°C) between 6°N and 12°N. The surface layer is nearly homogeneous
Figure 24 Vertical sections of temperature along sections WM8 (a) and SM5 (b) from west to east in the central Bay during winter and
summer.
Page 34
around 28°C with a thickness of about 50m north of 9°N. Further to the south the thickness increases considerably. Below the surface layer, the isotherms exhibit
spreading in the depth interval 50–200m between 8°N and 10°30′N. This features was noted by Colborn (1975) as important in the Andaman Sea. The spreading of
the isotherms indicates the presence of subsurface divergence, corroborating the reported occurrence of divergence at 9°30′N: 93°E during FebruaryMarch 1963
(Rao & Jayararman 1968a). Below 300m, the northsouth variation of temperature is small. In the depth interval 1100–1700m, vertical temperature gradients are small
(1°C drop over 600m), whereas in the Bay of Bengal the drop is 2.4°C for the same depth range during both winter and summer (see Fig. 23). This interesting feature
is considered to be attributable to the enclosed nature of the Andaman Seas.
Colborn (1975) studied the thermal structure of the Indian Ocean in a very detailed way using over 28000 observations (up to 1965). He divided the entire Indian
Ocean into 274 subareas horizontally and two depth domains vertically, viz., upper seasonal layer (0 to about 300m) and lower nonseasonal oceanic thermocline layer
(up to about 1200m). For the Bay
Figure 25 Vertical section of temperature along WM9 in the Andaman Sea during winter.
Page 35
of Bengal the data coverage was inadequate for some subareas. He found that variations in the thermal structure in the upper 500m are unique because of the
monsoons. He noted that in the Bay of Bengal, seasonal changes of SST resulted from the annual march of the sun and variations in cloudiness and wind speed. The
currents and river runoffs were found to influence the thermal structure considerably. He observed that the influence of high salinity water at intermediate depths on the
thermal structure was minimal owing to the remoteness of its source. Vertical heat transfer downwards from the surface and upwards from cold deeper layers was
found to maintain the thermal gradient layer at intermediate depths of the Bay of Bengal. In the Andaman Sea, the vertical thermal structure was influenced by vertical
motions of and within the permanent thermocline. Here also the important factors that control the annual thermal cycle were insolation, wind evaporation, river runoff
and currents.
Salinity fields
Surface
During the presummer monsoon (Fig. 26a) surface salinity varies between 32.4ppt in the northwestern Bay to 34.0ppt off the southern east coast of India. A tongue of
high salinity, aligned in a northeasterly direction, separates the low salinity waters of the northwestern and eastern Bay. Sewell (1929) showed a meridional variation of
surface salinity of about 2ppt with lower (32.0ppt) salinities in the northern Bay and northern Andaman Sea and higher (34.0ppt) salinities in the southern Bay during
MarchApril. Figure 26a also shows the same pattern of northsouth variation (see Fig. 29a for winter peak). Rao & Jayaraman (1968a) reported variation of surface
salinity between 33.37ppt and 34.23ppt along an 8°30′N section in the central Bay and between 32.36ppt and 33.6ppt along a 16°30′N section in the northwestern
Bay during MarchApril 1963.
During the summer monsoon, surface salinity decreases considerably to 21ppt at the head of the Bay (Fig. 27a) under the influence of increased river runoff from the
Ganges and Brahamputra rivers. An influx of fresh water of about 28.1×103m3s−1 flows into the Bay annually and about 50% of this comes during the summer
monsoon months (Table 1, see p. 7). High surface salinity values (34.0ppt) are found south of 15°N (around 81 °E). Low saline water (about 29.0ppt) spreads into the
interior of the Bay in a southwesterly direction from the head of the Bay and the northern Andaman Sea, These salinity tongues agree well with the circulation pattern in
Figure 42. Varkey (1986) also obtained a similar pattern of surface salinity distribution for the summer. Sewell (1929) shows a meridional variation of 4.5ppt in the Bay
of Bengal and 13.5ppt in the Andaman Sea during JuneAugust, He found lowest (20ppt) surface salinities in the northern Andaman Sea compared with those in the
northern Bay of Bengal (30ppt) and highest surface salinity in the southern Bay (34.5ppt) around 4°N.
During the postsummer monsoon, surface salinity shows a variation of 2.5ppt from 32.0ppt off the southern east coast of India to 34.5ppt in the southwestern Bay
(Fig. 28a). Low (33ppt) salinities are also seen west of the Andaman Islands. The low salinities at the southern east coast of India suggest the southwestward advection
of low salinity waters from the northwestern Bay with the weakening or withdrawal of the Indian Monsoon Current. Sewell (1929) showed highest meridional variation
of surface salinity during SeptemberNovember with lowest (20ppt) salinity in the northern Bay of Bengal and Andaman Sea and highest salinity in the southwestern
Bay. Relatively lower salinities of 26ppt were also reported at the mouths of the Rivers Godavari and Krishna.
Page 36
During the winter monsoon, surface salinity shows wide variations between 30.2ppt and 34.0ppt (Fig. 29a). Near river mouths, surface salinity is as low as 27.0ppt
(Fig. 6) off the central east coast of India and 31.4ppt in the northeastern Bay under the influence of freshwater influx from the Irrawady river. In the northwestern Bay,
salinities are between 33.0ppt and 32.4ppt, whereas Sankaranarayanan & Reddy (1968) reported surface salinities
Figure 26 Horizontal sections of salinity for presummer monsoon at 0m, 100m, 200m and 500m for Bay of Bengal.
Page 37
varying between 29.0ppt and 33.11ppt during January 1965 in this region, Suryanarayana et al. (1993) reported nearsurface salinity variation from 31.0 to 33.5ppt
during December 1983. These variations are within the possible range of fluctuations occuring as a result of changes in currents and river runoff. A salinity frontlike
pattern was noticed in the central Bay of Bengal with high (>34.0ppt) salinity waters in the south. This seems to be a conse
Figure 27 Horizontal sections of salinity for summer monsoon at 0m, 100m, 200m and 500m for Bay of Bengal.
Page 38
quence of the inward branching of the Northern Equatorial Current (Fig. 42, p. 58). Sewell (1929) found low salinities in the eastern Andaman Sea (see pp. 13–18 for
salinities off Thailand). Varkey (1986) found that the dilute surface water off the mouths of rivers along the central east coast of India was transported offshore in a
southeasterly direction by the northern part of the anticyclonic gyre in the southern Bay (Fig. 29a, 42). The large surface salinity variations observed during different
seasons are only a result of the wide fluctuations in river runoffs and surface currents. La Violette (1967) pointed out that the river runoff at the head of the Bay gets
mixed up and transported by the prevailing currents; towards the eastern parts of the Bay of Bengal and into the Andaman Sea if clockwise circulation prevails or
towards the east coast of India for counter clockwise circulation. Sasamal (1990) reported abnormal surface salinities (>35.2ppt) about 300km offshore along 19°30′
during FebruaryMarch. He explained this as due to local formation involving runoff and dry period.
100 metre surface

During the presummer monsoon (Fig. 26b), the salinity distribution is characterized by a large low salinity cell with a core salinity of 33.6ppt. A similar pattern of
isohalines is also observed at 50m and the increase of salinity from sea surface to 100m is 0.6ppt at the core. This cell extends westward up to the coast. During the
summer monsoon season (Fig. 27b), a tongue of high salinity with a core value of 35.2ppt, protrudes into the central Bay. Relatively large lateral gradients are noticed
to the east of the high salinity tongue as the salinity decreases towards the eastern Bay. Low salinities (<34.6ppt) are seen in the northwestern Bay and eastern Bay.
Sastry et al. (1985) and Murty et al. (1992b) identified this high salin
Figure 28 Horizontal sections of salinity for postsummer monsoon at 0m, 100m, 200m and 500m for Bay of Bengal.
Page 39
ity water as being due to the penetration of Arabian Sea high salinity water into the central Bay from the southwestern Bay at subsurface depths (75–100m).
During the postsummer season, a tongue of high salinity with a core value of 35.2ppt is present in the southwestern Bay (Fig. 28b) as a continuation from summer.
Outside the high salinity tongue, salinity decreases to 34.4ppt near the islands and 34.6ppt off Madras (13°N). Murty et al. (1992b) recorded a uniform value of salinity
(34.9ppt) at 100m in the
Figure 29 Horizontal sections of salinity for winter at 0m, 100m, 200m and 500m for Bay of Bengal (after Rao 1977).
Page 40
northwestern Bay during September 1983. During the winter monsoon (Fig. 29b), a tongue of high (35.2ppt) salinity penetrates into the Bay from the southwestern Bay
up to 7°N. This seems to result from the flow of Arabian Sea high salinity water into the Bay. Similar tongues of high salinity waters were seen during the two previous
seasons (Figs 27, 28). North of the high salinity tongue, salinity decreases in all directions, to, at its lowest,
Figure 30 Horizontal sections of salinity for the four seasons at 1000m (winter plot after Rao 1977).
Page 41
Figure 31 Vertical sections of salinity along WM7 (a) and SM4 (b) from head of Bay to equator during winter and summer.
34.0ppt in the Andaman Sea and at the head of the Bay. Considering the whole Bay of Bengal above 100m, it can be seen from Figures 26, 27, 28 and 29 that the
Bay of Bengal low salinity water, Indian Equatorial surface water (34.6–34.8ppt) and Arabian Sea high salinity water occupy the water body fully (see pp. 50–53).
The Arabian Sea high salinity water is only absent in the southern Bay during the premonsoon period.
200 metre surface

During the presummer monsoon season, salinity variation is marginal between 34.90ppt and 35.05ppt (Fig. 26c). The prominent low salinity cell noticed at 100m (Fig.
26b), seems to persist up to 200m, however, with very low lateral salinity gradients. The presence of Indian Equatorial Intermediate Water (Gallagher 1966) is noticed
intermittently around the low salinity cell (see Fig. 26d) for its presence throughout the Bay at 500m). During the summer monsoon season (Fig. 27c), salinity at 200m
varies between 34.95ppt and 35.10ppt and is higher by 0.05ppt than that of presummer monsoon salinities. Except for a single value of 34.95ppt off the central east
coast of India, salinities all over the Bay are over 35.0ppt and this indicates the stronger intrusion of Indian Equatorial Intermediate Water during summer than occurs
during the presummer monsoon season.
During the postsummer monsoon season (Fig. 28c), salinity variation is from 34.85 to 35.05ppt. During the winter monsoon, salinity at 200m varies between
34.9ppt and 35.0ppt (Fig. 29c). A large cell of low salinity (34.90ppt) occupies the central Bay. The pattern of isohalines in the southern Bay suggests a flow of the
North Equatorial Current advecting relatively high salinity water of the subsurface equatorial region.
500 metre surface

During the presummer monsoon, salinity varies between 35.02ppt and 35.05ppt (Fig. 26d). Variation of salinity during the summer monsoon is between 35.03ppt in
the northeastern Bay and 35.09ppt along 8°N and along 14°N, respectively (Fig. 27d). The distribution is characterized by zonallyorientated isohalines with relatively
high salinities (>35.06ppt) in
Page 42
Figure 32 Vertical sections of salinity along WM8 (a) and SM5 (b) from west to east in the central Bay during winter and summer.
Figure 33 Vertical section of salinity along WM9 in the Andaman Sea during winter.
the central Bay and southern Bay and low salinities (<35.06ppt) north of 16°N. During the postsummer monsoon season (Fig. 28d), salinity varies from 34.98ppt
west of the Nicobar Islands to 35.10ppt in the southern Bay and west central Bay. During the winter monsoon, salinity varies between 35.0ppt and 35.5ppt with the
high salinities in the northwestern Bay (Fig. 29d). Comparing all the distributions at 500m, it is seen that the penetration of the subsurface high salinity layer is maximal
during summer and postsummer periods. During summer and winter the whole Bay of Bengal is also occupied by this high saline layer.
Page 43

Figure 30 shows the seasonal distribution of salinity at 1000m. In general, salinity varies between 34.84ppt and 35.0ppt. During the presummer monsoon (Fig. 30a), a
cell of low salinity (34.94ppt) appears in the central Bay. However, lower salinities (34.92ppt) in the northwestern Bay and higher salinities (34.98ppt) off the southern
east coast of India are seen during the summer season (Fig. 30b). During the postsummer monsoon (Fig. 30c), salinities are much less compared with the summer
monsoon, with the lowest salinities (34.88ppt) near the Nicobar Islands. During the winter monsoon (Fig. 30d), the salinity variation is about 0.05ppt varying from
34.90 to 34.95ppt. It is interesting to see that the 1000m surface is almost free of the intermediate high saline core except for three observations during summer and
postsummer.
At 2000m (Fig. 22b, see p. 31) salinity varies between 34.78ppt and 34.84ppt and at 3000m (Fig. 22e, see p. 31) it varies by 0.06ppt between 34.72ppt and
34.78ppt. In general, relatively high salinities prevail in the southwestern Bay at 2000m. The salinity distribution at 2000m seems to indicate a general northwestward
movement of high salinity water from the southwestern Bay. This flow pattern is in agreement with the flow field deduced from geostrophic computations at these
depths (Zakliniski 1964, Wyrtki 1971, Murty 1990a, Murty et al. 1994). In the following three paragraphs, three vertical sections of salinity are presented for an
interrelated study of the horizontal charts.
Figures 31a and 32a represent typical meridional and zonal salinity sections along 91°E and 15°N, respectively, during the winter monsoon Along 91°E, two
pockets of low salinity centred at 8°N and 15°30′N are seen in the upper 50m. The salinity of the northern pocket is comparatively low (31.4ppt) and shows the
influence of freshwater influx from the Irrawady river towards the Bay of Bengal through the Preparis Channel. The depth of the halocline is 100m at 15°30′N, whereas
it is about 75m between 7°30′N and 9°30′E. The surfacing of the isohalines between 5°N and 7°30′N together with the low salinity pocket (33.2ppt) centred around
8°N is attributable to the North Equatorial Current advecting low salinity waters in the Bay of Bengal or Andaman Sea originating towards the west. The doming of the
isohalines around 18°N from 400m seems to be due to a cyclonic circulation cell (see Fig. 43, p. 59). The intermediate high salinity layer exhibits large depth and
thickness variations from south to north. Along 15°N (Fig. 32a), surface salinity decreases from 33.8ppt in the west to 31.6ppt in the east. Low salinity pockets
(<32.4ppt), influenced by the flow of fresh water from the Irrawady river, appear at 88°E and 93°E. Combined with the temperature distribution (Fig. 24a), it appears
that the fresh water is slightly warmer (about 0.5°C) north of the Andaman Islands. In the upper 100m, in the western part of the section, relatively high salinities are
observed in association with the northward flow (see Fig. 42, p. 58). The core salinity of the intermediate high salinity layer is higher (35.04ppt) in the western part than
in the eastern part.
Figures 31b and 32b show typical meridional and zonal distributions of salinity along 89°E and 13°N sections, respectively, during the summer monsoon. Along 89°
E low (<33.8ppt) salinity waters are seen north of 12°30′N with salinity decreasing northward to 32.6 ppt. Two cells of high (35.0–35.2ppt) salinities identified earlier
as the result of penetration of Arabian Sea High Salinity Water (Sastry et al. 1985, Murty et al. 1992) are located around 50m at 12°N and around 40m at 14°N. The
intermediate high salinity layer can be seen between 35.05ppt isohalines with its core (>35.1ppt) at 8°N. At deeper depths relatively low salinity waters are seen north
of 12°N between 500m and 1700m. Along the 13°N section (Fig. 32b), high surface salinities (33.6ppt) are seen off the Madras coast. Low
Page 44
salinities (32.4ppt) are observed at 91°30′E, because of the influx of fresh water from Irrawady river. The halocline is located at about 75m. Mild upward ascent of
isohalines is evident west of 84°E, suggesting a northward flow off the Madras coast (see Fig. 42, p. 58). The isohalines exhibit doming between 86°E and 88°E where
the high (35.0ppt) salinity cells appear at depths of about 90m due to the penetration of high salinity water mass. The intermediate high (>35.0ppt) salinity layer is
divided into large cells occupying the western and eastern regions, with relatively higher core salinity in the western cell.
Figure 33 represents the meridional distribution of salinity along 95°E in the Andaman Sea during the winter monsoon. Low surface salinities (32.0ppt) are seen here
owing to the runoff from the Irrawady river. Under the influence of this influx SST is also reduced slightly (see Fig. 25). The isohalines exhibit spreading centred around
9°30′N between 75m and 200m, and around 11°N between 100m and 300m suggesting subsurface divergences. In this subsurface divergence zone, the thickness of
the intermediate high salinity layer is about 350m, whereas it is about 400m at the southern and northern parts. The core salinity is slightly less in the northern Andaman
Sea.
Density fields and mixed layer
Surface
During the presummer monsoon season (Fig. 34a), low density (<20.5sigmat) waters occupy the east central Bay near the Andaman Islands and relatively dense
(21.0sigmat) waters are found along the coast and offshore areas. Rao & Jayaraman (1968a) noticed that during MarchApril 1963, the thickness of the mixed layer
was between 90m and 100m along 8°30′N, about 50m along 9°30′N in the region between the Andaman and Nicobar Islands and between 75m and 100m in the
western Bay.
During the summer monsoon season (Fig. 35a), the lowest (<13sigmat) density waters are seen in the northwestern Bay and the distribution pattern of density in this
area resembles that of the salinity. Density increases gradually towards the south and a tongue of low density water spreads southwestward in the central Bay. Under
the influence of strong southwesterly winds at the sea surface, spreading of diluted (due to a huge river runoff) surface waters takes place through the prevailing surface
currents and a number of eddies. South of 12°N, density increases to 22.0sigmat in the southern Bay, where relatively saline waters are found (Fig. 27a). The
thickness of the surface layer is effected mainly by windforcing over the Bay. The surface layer thickness decreased from 100m in the southern Bay towards the north
to around 20–30m along an axis orientated southwestnortheast (Murty et al. 1992b). Pockets of thick mixed layers coinciding with the anticyclonic circulation cells
were seen northwest of the Andaman Islands and off the Madras coast (Murty et al. 1992b). In the northwestern Bay, the thickness varied between 20m and 40m.
During the postsummer monsoon season (Fig. 36a), surface density varies between 20.0sigmat and 22.0sigmat. Flow of dense waters, characterized by high
salinity (Fig. 28a), from the southwestern Bay is seen up to 8°N. Low mixedlayer thickness is seen close to the coast under the influence of coastal up welling.
During the winter monsoon (Fig. 37a), density of the surface waters is less than 21.82sigmat (>600cl/t). Two cells of very low density are found off the central east
coast of India and northwest of the Andaman Islands where freshwater discharges from the Krishna and Irrawady rivers, respectively, enter the Bay. A large tongue of
high density water, protruding northwards, appears in the central part of the Bay between 5°N and 15°N
Page 45
and separates the two cells of low density water. Thus, strong lateral gradients occur in the eastern Bay and between 12°N and 15°N in the western Bay. Spreading of
low density water from the Andaman Sea towards the Bay of Bengal through the Ten Degree Channel is evident. The surface layer is thin (60m) between the equator
and 5°N and thick (>100m) at
Figure 34 Horizontal sections of density for presummer monsoon at 0m, 100m, 200m and 500m for Bay of Bengal.
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Figure 35 Horizontal sections of density for summer monsoon at 0m, 100m, 200m and 500m for Bay of Bengal.
8°N, while it is nearly uniform (about 75m) north of 10°N (see Fig. 23a). Distributions for the four seasons make it clear that the surface density variations within and
between seasons are controlled by insolation, evaporative cooling and influxes of saline or fresh water.
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Figure 36 Horizontal sections of density for postsummer monsoon at 0m, 100m, 200m and 500m for Bay of Bengal.
100 metre surface

During the presummer monsoon season (Fig. 34b) a large cell of low density water with a core density of 22.0sigmat is seen between 12°N and 17°N (see also Fig.
26b). The cellular structure suggests considerable geostrophic flow at this depth; a poleward flow in the western Bay and a zonal flow directed eastward along 17°N.
During the summer monsoon season (Fig. 35b), a broad band of high density water (>24.5sigmat) is seen in the central Bay. The density distribution indicates a
northward geostrophic flow in the eastern Bay and an anticyclonic circulation cell in the northwestern Bay.
During the postsummer monsoon season (Fig. 36b), density varies between 24.0sigmat and 25.0sigmat in the southern Bay of Bengal. During the winter monsoon
season (Fig. 37b), density varies between 24.96sigmat and 21.80sigmat. The northward increase of the isanosteres in the southwestern Bay indicates a northwesterly
flow of the North Equatorial Current. (See also Fig. 42). Cells of low and high density appear in the northern Bay indicating cyclonic and anticyclonic circulation cells.
200m surface
During the presummer monsoon season (Fig. 34c) the upper layer of low density cells persists up to this depth with a core density of 25.0sigmat. During the summer
monsoon season (Fig. 35c), density varies between 25.8sigmat and 26.4sigmat. The low density cell observed at 100m, west of the islands, still persists at 200m. In
the western Bay the distribution is characterized by alternate cells of low (26.0sigmat) and high (26.4sigmat) density waters.
During the postsummer monsoon season (Fig. 36c), density varies between 25.8sigmat
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Figure 37 Horizontal sections of density for winter monsoon at 0m, 100m, 200m and 500m for Bay of Bengal (after Rao 1977).
and 26.4sigmat. In the eastern Bay, density is uniform at 26.2sigmat. During the winter monsoon season (Fig. 37), density is low (high values of thermosteric
anomaly) in the northwestern Bay. Relatively high density waters are seen off the central east coast of India and the coast of Burma. The signature of the North
Equatorial Current in the southern Bay is also persistent at this depth.
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Figure 38 Horizontal sections of density for the four seasons at 1000m (winter plot after Rao 1977). See Figure 28 for sigmat
equivalents of cl/t.
500 metre surface

During the presummer monsoon season (Fig. 34d), density variation is between 27.0sigmat and 27.1sigmat, and two cells of high density appear in the northern and
western Bay.
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During the summer monsoon season also (Fig. 35d), density variation is between 27.0sigmat and 27.1sigmat. High density waters are noticed in small cells. In the
northwestern Bay a weak anticyclonic flow (opposite to surface flow) is seen (Fig. 44, see p. 60). During the postsummer monsoon season (Fig. 36d), density varies
between 27.0sigmat and 27.15sigmat. During the winter monsoon season (Fig. 37d), density varies between 27.07sigmat and 26.96sigmat.

The seasonal distribution of density at 1000m is presented in Figure 38. The density, in general, varies between 27.4sigmat and 27.5sigmat. Figure 22c, f (see p. 31)
presents distributions of density at 2000m and 3000m, respectively. As thermohaline variations are very small at these depths only standard deepwater features are
observed.
Water masses
Detailed studies of water masses of the Bay were nonexistent before IIOE. La Fond (1958) grouped the surface waters of the western Bay of Bengal into three
distinct classes, viz., the Northern Dilute Water (NDW), a transition water and the Southern Bay of Bengal Water (SBBW). NDW was considered to have been
formed as a result of large quantities of freshwater influx from this Ganges and Brahmaputra rivers. In terms of sigmat, density of the water is less than 19sigmat. The
transition water is a mixture of the NDW and the SBBW in the western Bay and has a density between 19sigmat and 21sigmat. The SBBW is the major source of
surface water in the Bay of Bengal with its density between 21sigmat and 22sigmat. However, it can be observed along the east coast of India as it is carried north by
the northeast coastal current. It is apparent that these results are limited to the Bay alone. Gallagher (1966) pointed out that the subsurface waters of the Bay of Bengal
are very uniform in all parts of the Bay north of 5°N and can be identified as the Indian Equatorial Intermediate Water (IEIW). Furthermore, he stated that this water
penetrates into the Bay at depths greater than 100–150m. In the central and southern Bay of Bengal, this water mass is present up to 1000m with its temperature
decreasing from 15°C to 5°C and the salinity remaining nearly constant (35.0 to 35.1ppt). He suggested that IEIW is formed as a result of mixing between high salinity
waters from the Arabian Sea and the North Indian Ocean Deep Water (NIODW). Rao & Jayaraman (1968b) identified Eastern Dilute Waters of IndoPacific origin in
the upper 100m and Indian Ocean Equatorial Water (T=4–16°C, S=34.8–35.2ppt) below 100–150m in the southern Bay of Bengal during FebruaryMarch.
Scherbinin (1969) studied the structure of water masses in the equatorial Indian Ocean. Sharma (1972) showed a salinity of 35.0ppt on 200cl/t isanosteric surface
appearing at 250m in the south central Bay of Bengal during the summer monsoon. This corresponds to the IEIW. Maslennikov (1973) noted that the surface waters of
the Bay of Bengal and the Andaman Sea could be regarded as a single type due to identical conditions of formation. He identified the penetration of waters from the
Arabian Sea into the Bay characterized by high salinity. It was found also that the vertical water structures in the Bay of Bengal and Andaman Sea are similar up to
700–800m.
Varkey (1986) studied in detail the water masses in the Bay during the winter. Figure 39 (a modified version from Varkey 1986) shows some typical TS diagrams
in selected areas (a) southwestern Bay (5–10°N and 81–89°E), (b) east equatorial Indian Ocean (92–97°E),
Page 51
(c) Andaman Sea (11–14°N and 95–97°E) and (d) northwestern Bay (13–16°N and 82–84 °E). From the figure it can be seen that the TS relations of the surface
waters of the southwestern Bay and east equatorial Indian ocean are more or less similar. Below the surface layer, waters of high salinity exceeding 35.0ppt occur
between 22.6sigmat and 27.4sigmat. The surface waters are cooler and fresher in the northwestern Bay (Fig. 39d) compared with those of the Andaman Sea (Fig.
39c). However, waters with salinity around 35.0ppt occur between 26.0sigmat and 27.04sigmat below the surface layer. At deeper depths, below 27.07sigmat, the
TS structures in the four areas are similar with an exponential decrease. The subsurface saline (>35.0ppt) layer was identified (Varkey 1986) as Intermediate High
Salinity Water (IHSW) of the Bay of Bengal and represents the above IEIW wherein the Persian Gulf Water (PGW) and Red Sea Water (RSW) enter the Bay.
Varkey (1986) identified the low salinity surface water as the Bay of Bengal Low Salinity Water (BBLSW). Water Mass E (suggested by Rochford 1964) and the
Arabian Sea High Salinity Water (ASHSW), are also present in the subsurface layer during the winter monsoon. The BBLSW appears in the northern Bay and
Andaman Sea and is formed as a result of great dilution and corresponds to the salinity domains marked as the NOW and transition water by La Fond (1958). Both
the Water Mass E and the ASHSW are found to overlap each other in the TS plots (see Fig. 39a). While the ASHSW is observed only in the southern Bay and off
Sri Lanka during the winter monsoon, it penetrates into the Bay at depths of 50–100m in the open sea areas of the Bay of Bengal between 4°N and 14°N during the
summer monsoon (Sastry et al. 1985, Murty et al. 1992b).
Varkey (1986) also noticed that the subsurface layer around the pycnocline is very diluted in the northern Bay, western Bay and northern Andaman Sea regions
(Figs 39c, d). He assigned thermohaline indices for the BBLSW as T=27°C, S=33.0ppt around 21.2sigmat in the northern Bay. It was also pointed out that this
BBLSW is less homogeneous in its TS values compared with the intermediate and deep water masses because of the wide range of environmental conditions
experienced at the surface, such as runoff, precipitation, evaporation and high salinity currents. The thermohaline indices for the IHSW core are assigned as T=9.6°C,
S=35.03ppt at 27.07sigmat around 500m. In the inner subsurface regions of the Bay (outside the coastal Bay), a hypothetical water with thermohaline indices T=27°
C,
Figure 39 Typical TS diagrams for areas (a) southwestern bay (b) east equatorial Indian Ocean (c) Andaman Sea and (d)
northwestern Bay.
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S=34.6ppt (around point C in Fig. 39d) is envisaged as a mixture of BBLSW at the surface and IHSW at intermediate depths. Varkey (1986) also found turbulent
mixing coefficients to be very high at the pycnocline compared with the mixed layer (see pp. 53–56). In the pycnocline region, even although high static stability is
observed, high current shear and internal wave breaking (resulting in high eddy coefficients) cause vertical mixing across the pycnocline. This is one of the ways by
which nutrientrich subsurface waters are brought to the surface in the Bay.
Wyrtki (1961), Rochford (1964) and Suryanarayana et al. (1993) have identified the ASHSW at subsurface depths (50–100m) in the southern Bay towards the
equator. From Figures 31a and 32a, it can be seen that ASHSW does not appear in the central Bay (15°N) and eastern Bay (91°E) during the winter monsoon.
However, its penetration in the depth range of 75–125m can be seen between 12–14°N and 87–89°E during the summer monsoon (Figs 31b, 32b) and can be
influenced by the surface divergences and gyres. Wyrtki (1961, 1971), Rochford (1964), Rao & Jayaraman (1968a) and Varadachari et al. (1968b) also noticed a
high salinity (>35.0ppt) water mass in the intermediate depth range of the Bay of Bengal and attributed this to penetration of saline PGW and RSW at different depths
greater than 200m during different seasons. Murty et al. (1992) and Suryanarayana et al. (1993) observed a layer of high salinity water (35.0–35.1ppt) in the depth
interval of 200–900m between 26.0sigmat and 27.4sigmat isopycnals with its salinity decreasing marginally towards the northern Bay. It has been suggested that this
intermediate nearly isohaline layer was the result of penetration of a similar isohaline layer (35.7ppt) in the depth interval 250–800m, which formed along the Arabian
coast in the Arabian Sea and was itself the result of high salinity mixing of PGW and RSW (Premchand 1981, Sastry et al. 1985, Premchand et al. 1986). Premchand
et al. (1986) pointed out that the salinity of the isohaline layer in the Arabian Sea progressively decreases as it moves away from the Arabian Sea to the eastern Indian
Ocean because of its subsequent mixing with ambient waters along its path; these processes could give rise to the observed TS structure in the Bay of Bengal. The
thickness of the IHSW layer fluctuates in the western Bay (Figs 31a, 32a). During the summer monsoon, the salinity of the IHSW is high and reaches up to 35.1ppt at
8°N, 89°E; this water appears in large cells in the western and eastern Bay (Figs 31b, 32b), and indicates the seasonal variation in thickness and salinity of the high
salinity layer particularly in the eastern Bay (Figs 32a, 32b). This IHSW is also noticed in the Andaman Sea during the winter monsoon (Fig. 33) although its thickness
and salinity show variations due to zones of divergence (central region) and convergence (northern and southern regions). The maximum salinity of this layer is 35.06ppt
in the southern region (Fig. 33).
At depths greater than 1500m, Northern Indian Deep Water spreads into the Bay of Bengal. This water mass is formed by mixing Arabian Sea Intermediate Water
and Indian Ocean Bottom Water (Gallagher 1966). The deep water has a temperature between 1.6°C and 2.8°C, a salinity of 34.68 to 34.78ppt and an oxygen
content of 3.18mll−1 to 4.17mll−1 (Gallagher 1966). Wyrtki (1971) showed that between 2000m and 3000m salinities and potential temperatures vary between
34.75ppt and 34.74ppt and 2.6°C and 1.4°C, respectively (see also Fig. 22). However, in the Andaman Sea basin, the deep water has a nearly uniform temperature
of around 4.8°C and salinity of 34.9ppt from the sill depth (around 1300–1500m) to the bottom because of its enclosed nature. Ramesh Babu & Sastry (1976)
compared the deep and bottom water characteristics in the Andaman Sea and Bay of Bengal and found that potential temperatures in the Andaman Sea varied within a
narrow range, 4.63–4.85°C between 1894m and 3088m, in contrast to the values in the Bay of Bengal where the potential temperature decreased with depth (2.17°C
at 2240m and 1.48°C at
Page 53
3070m). The salinity was nearly uniform (34.8ppt) in the entire water column of the Andaman Sea below 1300m whereas in the Bay of Bengal the bottom salinity
decreased to 34.74 ppt. Sewell (1932) showed that the eastern drift of the Antarctic bottom water was first northwards between Carpenter’s Ridge and the Andaman
Nicobar Ridge; it then established a rotary movement (cyclonic flow) in the central Bay of Bengal. The western branch of the bottom drift moves towards the east coast
of India and then northeastward up to the head of the Bay of Bengal and forms a clockwise flow in the eastern Bay. Bennette (1970) stated that “the deep water
structure of the northern Indian ocean (1700–3000m) is maintained in the same way of that of the deep pacific ocean….” This similarity he attributed to the global
northward flow of the Antarctic Bottom Water in all oceans and its subsequent ascending to the intermediate depths (up to 600m in the Arabian Sea).
Turbulent mixing and vertical advection

Studies relating to turbulent mixing in the Bay, both vertical and horizontal are very few. Available values of eddy coefficients (Kz and Kh ) are indirect estimates from
wind thrust and general hydrographic data. However, these estimates do give a vague picture of the situation in the Bay. Estimates of eddy coefficients in the literature
show a very wide scatter depending upon sensors and instruments, processing methods, and areas of study like surface mixed layer, thermocline, midocean, fjords,
estuaries, etc. Sverdrup et al. (1962) in their summary of eddy viscosity from different oceanic regions using different techniques, show a range from 0 to
7500gcm−1s−1. Pollard (1977) found a range of 20 to 500cm2s−1 in the surface layers of different environments.
Varkey (1986) studied the seasonal (winter) mean thermocline layer (0–400 m) and annual mean logarithmic layer (400–1200m) in the Bay in detail and evaluated a
vertical profile of Kz (Fig. 40). His mean values are as follows: K400=8cm2s−1, Kgoo= 16cm2s−1 and K1200=24cm2s−1. (The corresponding mean vertical velocities
are, W400=4.9×10−5cms−1, W800=9.8×10−5cms−1 and W1200=14.7×10−5cms−1). The values presented in Figure 40 (for the area 93–95°E and 4.5–6.5°N) can
change from season to season and area to area. But, for the same season the relative variations can also be considered to remain the same for other areas of the Bay.
(The corresponding mean vertical velocity (W) profile is shown in Figure 41). The most conspicuous feature of the seasonal mean Kz profile is the high values at the
upper half (75–100m) of the pycnocline; about 100 times the value at 225m and about three times that at surface. This high value at the pycnocline can ensure large
vertical eddy transport irrespective of the high density stratification in the Bay because of abundant freshwater inflow from rivers (see pp. 7–8 and 19). This high Kz
value was found to be due to the high current shear just above the main pycnocline and the resulting instability and internal wave breaking in the Bay (Varkey 1986).
Also shown in Figure 40 are values for transition layer (c. 1600m) and deep layers in the Bay (1750–2250m) and in the north Indian Ocean (1700–3000m). The
deep layer value (7cm2s−1) for the Bay was computed (Varkey 1986) using a mean W/Kz value of to 1.13×10−4cms−1. Bennette (1970) calculated a value of
2.5cm2s−1 for the deep north 1.61×10−5cm−1 for the full Bay excluding the Andaman Sea and a mean value of W2300 equal Indian Ocean, and the above value of
7cm2s−1 for the Bay is about 3 times this. The transition layer (1350–1750m) value (Fig. 40) was computed using a mean value of W/Kz as 7.6×10−6cm−1 and W2300
as mentioned above and serves as an approximation only. This is not strictly correct since the logarithmic approximation is not valid in this layer because of
Page 54
Figure 40 Vertical profile of mean KZ in the northeastern Indian Ocean.
the dominant lateral mixing (Bennette 1970). Varkey (1986) also found that horizontal flows are relatively high in this transition layer.
There exist many estimates of Kh reported by various researchers in different parts of world oceans. Sverdrup et al. (1962) summarized estimates of Kh and found
that they vary between 1×108gcm−1s−1 (for 400–2500m range) and 2×106gcm−1s−1 (for 200–400m range) through 4×107gcm−1s−1 (0–200m range). Defant (1961)
found a value of 5.5×107cm2s−1 for a Mediterranean water tongue that flows into the Atlantic. Garrett (1979) in his review found a wide range of 102 to 104 for Kh
estimated by various workers. Kh values also cover a very wide range and, like Kz, different sensors, various processing methods, and widely different environments
contribute to the discrepancies.
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Figure 41 Vertical profile of mean Kh and W in the thermocline layer of Bay of Bengal.
The values of Kh presented in Figure 41 can vary several times as a result of using crude methods (Varkey 1986). But this is of no significance considering the wide
range (105 to 108) of values within the range 20–350m found in the Bay. The relative profile emphasizes some noteworthy features. Kh values show a conspicuous
increase (>108) just below the main pycnocline (75–125m) or over the lower half of the thermocline. Bennette (1970) estimated Kh in the northern Indian Ocean from
Red Sea salinity maximum as 7×107cm2s−1 for bimonthly horizontal distribution and as 3×108cm2s−1 for mean annual horizontal distribution.
Grand mean vertical velocities computed (Varkey 1986) for a square in the southern Bay (4.5°–6.5°N and 93°–95°E), using the same methods used for Kh
computations, are also shown in Figure 41. The W values (upwelling) progressively increase from 4×10−5cms−1 at 350m to c.0.6cms−1 at the surface. Individual
vertical profiles at subgrid points within the square showed upwelling and downwelling layers (Varkey 1986). The sudden increase of Kz and Kh within the
pycnocline, together with the regular increase of W in the thermocline, suggests again that the exchanges across the pycnocline are controlled by turbulent processes.
Babu (1987) computed monthly vertical velocities at the base of the surface Ekman layer using monthly mean wind data (Hastenrath & Lamb 1979) for 1° squares
in the Bay excluding the Andaman Sea. The W values for the summer monsoon months of June, July, August and September lie within the range +16×10−4 to
−8×10−4cms−1 with upwelling (positive) and downwelling (negative) areas. For December and January, peak winter monsoon months, W values range between
+6×10−4cms−1 and −8×10−4cms−1 with sinking all along the northwest coast from Madras to Paradip. During MarchApril, the W values are found to be low
(<2×10−4 to −2×10−4cms−1) suggesting that upwelling and sinking in the Bay is pri
Page 56
marily windinduced. Babu et al. (1989) also computed W values for August September from instantaneously observed winds and found values ranging between
+2.0×1 0−3cms−1 to to −2.5×10−3cms−1. The differences seem to be due to differences in types of data. Even although all these estimates are at the base of the Ekman
layer, the fact that this layer varies in depth from season to season depending upon wind velocity should be given due consideration.
Comparatively, the W values obtained by Varkey (1986) and Babu (1987), using mean seasonal hydrographic data and mean monthly wind stress show
respectively, good agreement at the base of the Ekman layers for the winter monsoon period (0.4×10−4cms−1 and 10−4cms−1). These estimates also agree well with
those of Veronis (1969), where a value of 0.46×10−4cms−1 was obtained at the base depth of the thermocline. Comparisons for other seasons are not possible since
Varkey (1986) studied only the winter monsoon period. For the deep Bay (2300m) Varkey (1986) calculated a value of W equal to 1.1×10−4s−1 assuming that a net
outflow of 60×1012m3yr−1 takes place across 6°N between the surface and 1200m and that this water uniformly ascends from the deep Bay. The W values evaluated
from mean yearly temperature profiles of all 5° squares of the Bay together (see pp. 53–54), are valid only in the layer 400 to 1200m and assume a total and only
balance between vertical advection and turbulent diffusion. Here, results of another study (Bahulayan & Varadachari 1985) in the equatorial Arabian Sea between 50°
E and 60°E, using data for winter (February 1967), are of comparative significance. They obtained W10 as 0.1cms−1 and W50 as 0.05cms−1 whereas Varkey (1986)
got a value of 0.6cms−1 at 10m. (But their methods differed widely, since Varky (1986) used depth dependant K2 and Kh ).
Currents and water transports
Currents
All atlases and charts show varying current patterns in the Bay (Anonymous 1953, 1960, Cutler & Swallow 1984 etc.). Poornachandra Rao (1956) found, from
geostrophic computation, along a 185km long section off Visakhatpatnam, that during March (a) the surface current was strongest (c.41 cms−1) and northeasterly at
75km offshore, (b) currents decreased gradually with depth up to about 300m where velocity c.7cms−1 and (c) belts of strong and weak flows were present across the
section. Balaramamurty & Ramasastry (1957) found northeasterly flows between Visakhapatman and Kakinada during DecemberFebruary, typical winter monsoon
months. During the same period from the Bangladesh coast to Mahanadimouth the currents followed a downthecoast direction. During MarchApril, they found very
small density fluctuations from Visakhapatnam to Madras and inferred the surface currents to be weak and variable. During OctoberNovember, postsummer
monsoon months, the currents flowed down along the same coast over a belt about 100km wide. La Fond (1957) also observed at times very high velocities of 5 knots
of Sacramanto shoals near Godavari mouth. This could have been attributable to river flow and tidal effects. La Fond (1958) found that, along the central east coast,
north of Waltair (about 10km up the coast from Visakhapatnam), the surface currents were northeasterly during March with speeds from 0.5 to 1.7 knots. Near to the
coasts during October 1952, currents were to the southwest with a maximum velocity of 2.5 knots.
Nejman (1963) found a clockwise gyre at the surface for JulyOctober in the southern
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Bay, influenced by the North Equatorial Current. At 1000m the circulation changed to an anticlockwise direction. In the south Andaman Sea, also, a clockwise cell was
noted at the surface and at 50m. Fomicev (1964) found from dynamic computations that circulation in the northern Bay (north of 12°N) during February, a typical
winter month, consisted of four cells, one clockwise and three anticlockwise, including one in the Gulf of Martaban, with surface velocities of c.50cms−1. The southern
Bay contained a clockwise fullBay gyre with its southern part joining the North Equatorial Current. During March, the deep Andaman Sea contained a double gyre
with a clockwise southeastern cell and an anticlockwise northwestern cell. At 500m these cells merged to form one gyre. Varadachari & Sharma (1967) found, from an
analysis of streamlines using observations from various ships, that in the southern Bay (south of c.12°N) there existed seasonallyreversing currents as parts of the west
flowing North Equatorial Current during and as part of the eastflowing Indian Monsoon Current during summer. Varadachari et al. (1968a) from geostrophic
computation for June and July found two clockwise cells centred around 11°N and 83°E and 16°N and 85°E in the southwestern Bay with an enveloping coastal
current. Velocities varied from approximately 3knots at the surface to 0.1knot at 400m. La Fond & La Fond (1968) suggested that clockwise eddies are a seasonal
feature of the Bay. They described two eddies, one about 240km in diameter in the northeastern Bay by about 320km off the mouth of the Ganges and the second
(c.540km in diameter) between Madras and Visakhapatman. Rao & Jayaraman (1968a) found zones of convergences and divergences at the surface and at
subsurface depths in the southern and central Bay during FebruaryMarch. They found a shallow (<30m) southwesterly flow of low salinity water across 15°N
between 91°E and 94°E and attributed this to flow of fresh water from the Irrawady river and the prevailing northnortheast winds. Düing (1970) divided the year into
five characteristic time periods for studying the dynamic topographies of the sea surface (north of 20°S) as winter, spring, early and late summer and fall. He observed a
complex pattern of highs and lows in the topographic charts for the various periods and found their numbers to be more in the transition periods. He also presented a
theoretical study investigating the circulation pattern of the north Indian Ocean.
Shcherbinin (1973) computed geostrophic currents for the Indian Ocean (north of 10°S) for both winter and summer and found substantial changes in the circulation
patterns between the two seasons. But the drastic seasonal changes were limited to the Bay and Andaman Sea up to about 300m. During winter the North Equatorial
Current flows west from Sumatra to Sri Lanka where it branches into two, one running north along the east coast of India to form part of the anticyclonic gyre of the
Bay. During summer an anticyclonic circulation existed between the east coast, 87°E, and <16°N. At the head of the Bay one weak cyclonic cell was noted. Another
anticyclonic cell was seen at the southern Andaman Sea. This summer circulation pattern persisted to about 100m in the Bay. Maslennikov (1973) suggested velocities
between 0.3knots and 1.3knots for the northern and southern sides of the anticlockwise cell in the Gulf of Martaban. This cell, he attributed to southwestward slope
currents and inflow from the Bay into the northern Andaman Sea. Krishna Rao (1974) observed from satellite pictures that during September the surface circulation
between Visakhapatnam and c.19°N was anticlockwise along the coast while further south this current turned southeast. He also noted that between 15°N and 17°N
the coastal circulation was towards the north turning eastwards at the northern limit, making the region between 17°N and 18°N a confluence area. A similar frontal
region was observed by Rao et al. (1987) between the Godavari and the Mahanadi, separating two regimes of coastal current during JuneJuly. Ramesh Babu &
Sastry (1976) observed that the flow in the Andaman Sea consisted of cyclonic and anticyclonic cells of 100–200km
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extending up to depths of 100–150m. Kolla et al. (1976) observed comparatively strong bottom currents in a broad valley in the deep (c.4000m) western Bay
between 5°N and 12°N. Sea floor photographs of some objects such as faecal pellets suggested southward currents. One direct measurement at the bottom showed a
velocity of 7cms−1 towards the south. Gopalakrishna & Sastry (1985) found that north of Visakhapatnam strong southwesterly flows existed up to north of the mouth
of the Mahanadi during August September. Sarma & Rao (1986) using moored current meters, observed hourly mean speeds of 27.2cms−1, 18.4cms−1 and
18.2cms−1 at 15 m, 115 m, and 240 m, respectively, at a station just outside the shelf (depth of c.250m) off the mouth of the Godavari during September.
Corresponding maximum speeds were 45.7cms−1, 40.7cms−1 and 34cms−1. At all depths directions varied, flowing 180° out of phase within 24 to 30h. At 15m
directions varied between SW, SE and NE.
Varkey (1986), after a detailed study of the oceanography of the Bay and Andaman Sea, suggested a threegyred mean seasonal circulation pattern for the two
monsoons; (a) a northwestern gyre (G1) between coastline, c.13°N and c.89°E, clockwise during winter and anticlockwise in summer, (b) a southern gyre (G2) in the
area south of c.13°N, clockwise during both winter and summer and (c) a double gyre (G3) in the Andaman Sea with clockwise and anticlockwise flows for both
seasons. During winter, G1 was found to reverse its flow at 500m and maintain the same pattern at 1000m. G2 and G3 were found to maintain their surface directions
at 500m and 1000m during winter (see Figs 42 & 43 for slightly modified patterns). He pointed out that couplings between the gyres were affected by eddies of
various sizes and directions.
From satellite imagery for February 1985, presented by Legeckis (1987), a clear branching of the North Equatorial Current around 5°N of Sri Lanka can be seen.
The northern branch flows northwards following the coast as a “western boundary current” up to about 19°N. His plates clearly show a flow pattern similar to G1 and
G2, the bifurcation of this northern branch into two clockwise gyres occurring at about 13°N. Legeckis also found evidence for a number of eddies of different sizes
and shapes broken off from the main streams,
Figure 42 Geostrophic current marks transient cells.

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Figure 43 Geostrophic current vectors marks transient cells.
drifting with different velocities (e.g. 45cms−1, 30cms−1). He also noted that the “western boundary current” was variable or unstable depending on the North
Equatorial Current. He observed a flow of cold water southwards along 92°E up to 14°N, a branch of which turned westwards between 16°N and 18°N. This cold
flow corresponds with that reported by Rao & Jayaraman (1968a). Suryanarayana (1988) worked out coastal currents from 10° to 20°N over a coastal belt of about
250km width for winter and summer. He also found the system to consist of a few cyclonic and anticyclonic cells at the surface. In the vertical section also, the flows
showed zonations (or bands) with southwesterly and northeasterly directions down to bottom over the shelf. Molinari et al. (1990) observed, from satellite tracked
drifting buoy data, a clockwise gyre in the northwestern Bay during FebruaryMarch similar to G1 in winter. Khokiattiwong (1991) found that off the west coast of
Thailand (between 6°N and 7°40′N) during the winter monsoon, shallow water currents were strongly influenced by tidal currents. Away from the coast (>25km),
unaffected by the coastal islands, currents were stronger and tended to flow northwards. During MarchApril, Shetye et al. (1993b) found a strong coastal current
flowing northeastwards from 10 to 20°N. From 14°N onwards this current shifted offshore by c.110km with a more easterly component and formed the western
branch of an anticyclonic gyre extending to about 20°N and 90°E. It was also noted (Shetye et al. 1993b) that this gyre intensifies from January (3×106m3s−1) to
MarchApril (5×106m3s−1) and by June it disintegrates. These gyres, reported for the winter monsoon by various workers, agree well with G1 and G2 of winter as
described by Varkey (1986). (Here it is noteworthy that Wyrtki’s atlas (IIOE) shows an intensification of the circulation in the northwestern Bay during MarchApril
from that during JanuaryFebruary). M.T.Babu et al. (pers. comm.) also observed a meandering “western boundary current” between 13°N and 18°N during March
April 1987. From 13°N to 18°N it almost doubled its velocity to 1.0ms−1. This current left the coast at 17.5°N turning eastwards to form the northern part of an
anticyclonic gyre. Two coldcore cyclonic eddies were also observed by them at 14°N (which corresponds to TC3 in Fig. 42) and 18.5°N near the coast.
Suryanarayana et al.
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Figure 44 Dynamic topographies of different surfaces during summer.
(1993) identified a poleward flowing current west of the AndamanNicobar Islands during OctoberNovember 1983. (This seems to correspond to TC1 in Fig. 42).
Murty (1990) studied currents and transports in the Bay excluding the Andaman Sea during summer. The surface currents relative to 1000db agreed with the
patterns presented in Figure 42a. At 500m surface (relative to 1000db), G1 reverses its direction and becomes a clockwise flow (Fig. 44a). Near to the coast around
15°N a small southward current flows. This seems to be a transient cell coupling G1 and G2. The southern gyre G2 exists at 500m also, in the same way as at the
surface. Similar to the TC2 in winter at 500m (Fig. 43a), a transient cell also exists during summer in the same area. At 1000m (Fig. 44b), relative to 2500db, G1 exists
in the same way as at 500m. G2 shifts slightly towards the northwest and is a closed flow. Coupling between G1 and G2 is effected by an anticlockwise eddy and a
southward coastal flow.
During July 1984 Babu et al. (1991) observed a cyclonic circulation pattern of elliptical shape (400×200km) above a depth of 300m around 17°40′N and 85°19′E
with the major axis parallel to the coast. During JulyAugust, Shetye et al. (1991) found a cyclonic gyre centred around c. 17°N and c.85°30′E parallel to the central
east coast between 14°N and 19°N stretching to about 400km offshore. These compare well with G1 for the summer season.
In some of the above works different levels of no motion were used for the geostrophic computations, e.g., 500, 1000, 3000db. Results obtained using 500db as
reference level were reviewed cautiously taking into consideration its inadequacy to bring out the real currents at the surface. For other works wherein levels of no
motion were 1000db or >1000db, the results were considered undistorted (see Wyrtki 1971, Varkey 1988 for more details on reference levels in the northern Indian
Ocean).
Modelling studies carried out in the Bay of Bengal and Andaman Sea also showed the existence of three steady, seasonal gyres. Potemra et al. (1991) studied the
circulation in the northeastern Indian Ocean and the surrounding equatorial area up to c. 10°S using a multilayer numerical model driven by monthly mean winds. During
February and March the circulation pattern consisted of three gyres; (a) a clockwise gyre (G1) in the northwestern Bay with a strong western boundary current, (b) a
weaker clockwise gyre (G2) in the southern Bay and (c) a double (with clockwise and anticlockwise parts) or single gyre (G3) in the
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Andaman Sea. During summer (June and August) the surface circulation also consisted of three gyres; (a) an anticlockwise gyre (G1) in the northwestern Bay, (b) the
northern part of a clockwise gyre (G2) in the southern Bay and (c) an anticlockwise or clockwise gyre (G3) in the Andaman Sea. The steadiness and strength of the
gyres and other currents in the Bay seem to depend on the development and shift of the North Equatorial Current. The bifurcation, sense of rotation and strength of the
currents in the Andaman Sea also seem to depend on the North Equatorial Current and northern closing of the AndamanNicobar chain of islands in the model. Yu et
al. (1991), through their numerical experiments, show that the seasonal reversal (cyclonic in summer to anticyclonic in winter) of G1 in the northwestern Bay is caused
partly by a remote forcing due to monsoon winds in the equatorial Indian ocean. They also observed a twogyred system during transition periods for the Bay area
alone. Unnikrishnan & Bahulayan (1991) and Bahulayan & Unnikrishnan (1992) studied the circulation in the Bay of Bengal and Andaman Sea using a vertically
integrated numerical model for the two seasons and intervening periods. They found two anticyclonic gyres between 13°N and 18°N and 6°N and 12°N (G2) during
June and July. A transient circulation cell was also noted between the coast and 18°N, which reversed from the anticyclonic to the cyclonic between June and July, i.e.
within the same season. During December, they obtained an anticyclonic gyre (G1) in the northwestern Bay and a transient anticyclonic cell in the head of the Bay
(c.200km offshore). The big cyclonic gyre in the area of G2 seems to be erroneous in direction and strength because of the inadequacy of boundary conditions. The
directions of G1 during June and July are also inconsistent with all other results because of the same defect. During September and October (postsummer monsoon
transition period), currents were very weak and variable and during April currents were weak and formed into three transient anticyclonic cells in the area north of 12°
N.
The early results before IIOE (1962), limited to a narrow belt along the east coast, clearly brought out the seasonality, depth dependence, zonations, and frontal
nature of the currents. Some of these works were based purely on surface winds and hydrographic observations. Some observations could have been strongly
influenced by river flow and tidal currents (e.g. La Fond 1957). Results published during and after IIOE, beginning with Nejman (1963), showed that the circulation in
the deep Bay of Bengal and Andaman Sea consists of well developed, strong, seasonal gyres, transient and/or unsteady current cells, and transient migratory eddies.
The gyres seem to be of high speeds compared with the other transient currents. The presence of various gyres, and other transient currents, seem to give rise to a
number of frontal areas, both horizontally and vertically. A close analysis of these results shows that the factors that cause, change, and shift the various currents are: (a)
wind thrust from seasonal winds, (b) river runoff from the KrishnaGodavari, MahanadiGangesBrahmaputra and IrrawadySalween systems, (c) the Andaman
Nicobar chain of islands, and (d) the North Equatorial Current system. The threegyred mean seasonal circulation patterns with transient migratory eddies, as suggested
by Varkey (1986), are well supported by later works using satellite imagery, temperaturesalinity data, and numerical models. Results from Potemra et al. (1991)
demonstrated the ability of a model to simulate the seasonal and depthwise changes of currents in the area. This model could have simulated the current system much
more realistically if the huge runoffs (see Table 1) from the northern coasts and the Preparis channel were suitably incorporated. The controlling effect of the North
Equatorial Current and Indian Monsoon Current on the circulation in the Bay is well demonstrated by Yu et al. (1991). A comparison of the results of the above two
works with those of Bahulayan & Unnikrishnan (1992) brings out the inefficiency of the southern boundary condition used by the latter authors, resulting in changes of
direction of G1 and G2 dur
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ing summer and winter, respectively.

Modified mean threegyred circulation patterns for the two seasons at the surface and at 500 and 1000m for winter (only) are shown in Figures 42 and 43. In
addition to the gyres, three transient cells are also shown for the winter, the rotation, speeds, sizes, and locations of which can vary depending on the gyres and other
controlling factors mentioned above. The effect of transients as a result of the runoff from KrishnaGodavari on G1 gets masked by the strong “western boundary
current”. Hence transient cells like TC1 and TC2 are not clearly observed here. The directions and speeds of the gyres in the Andaman Sea during winter and summer
would be, in reality, variable around those shown in the figures, whereas in the Bay of Bengal the mean directions of the G1 and G2 would always be the same with
speeds depending on factors mentioned earlier.
High geostrophic surface velocities (c.50cms−1) are observed (Fig. 42) along the coastal side of G1, northern and southern sides of G2 and the western side of G3
during the winter. Molinari et al. (1990) got 50 to 62cms−1 at G1 during February and March from drifting buoys. At 500m the mean velocities are c.12cms−1 over all
the Bay of Bengal and Andaman Sea (Fig. 43) except along the coastal side of G1 (c.30cms−1). At 1000m also (Fig. 43) the coastal side of G1 shows high velocity
(c.17cms−1) whereas in other areas velocities are c.10cms−1. The monthly mean surface velocities for G2 (at 7°N and 85°E) during June was found to be 62cms−1 and
for G1 (near to the coast) during September 75cms−1 (Molinari et al. 1990) from drifting buoys. The observed and computed values of surface currents fall within the
range of 1 to 1.5 knots including both seasons.
Transport across different sections

Varkey (1986) worked out mean seasonal (summer and winter) and mean yearly water transports, across 6°N section joining Sri Lanka and Sumatra, for various
layers, e.g. surface to 23sigmat, 23 to 26sigmat, 27.3 to 27.5sigmat, up to 1250m (Fig. 45). He found that during summer the net (inflowoutflow) transport is into
the Bay from the surface to 1000m.
Figure 45 Mean seasonal and mean yearly water transports across 6°N; a, winter, b, summer and c, yearly. shows net
transports.
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Figure 46 Water transports across various sections from surface to 2500m in the deep Bay. Enclosed numbers are net sectional
transports with direction.
During the winter two layers were observed, viz., surface to c.600m and c.600 to 1250m with net transports outwards and inwards, respectively. Yearly mean
transports showed threelayered structures, viz., surface to c.100m, c.100 to c.600m and c.600 to 1250m with net transports inwards, outwards and inwards,
respectively. The net transports for mean winter, summer and full year from surface to 1250m are 26.4×1012m3 per month (outflow),
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42.9×1012m3 per month (inflow) and 5×1012m3 per month (outflow). Hence, the net surface exchange volume for E+P+R (see pp. 9–10) are only 0.4% and 1.4% of
the oceanic transports during winter and summer, respectively, considering the full Bay north of 6°N.
Murty (1990a) computed currents and transports across various zonal (4 to 19.5°N at 2° intervals) and meridional (85°E, 88°E and 91°E) sections of varying length
for the summer in the Bay of Bengal. The results showed welldefined zonations (vertical bands) of opposing flows from surface to large depths (>1000m) both across
zonal and meridonal sections. It was found that north of 12°N the Bay has a net outflow and south of 12°N a net inflow for the depth range 0 to 2500m (Fig. 46). This
seems to be in agreement with the horizontal currents’ zonations (Figs 42, 43) wherein the area 12–13°N is found to be of a frontal nature. Across 12°N (from 80.5 to
92°E) the upper 1000m layer showed a net outflow and the lower layer (1000 to 2500m) a net large inflow. Rao & Murty (1992) computed transports across various
sections in the Bay for winter and found that transports are directed northwards across all sections (6 to 20°N at 2° intervals) except across 16°N and 20°N. This is
also in agreement with the gyral current pattern between c.15°N and c.20°N forming frontal regions around 12–13°N and 20 °N and the well developed northeasterly
flow all along the western boundary of the Bay. Considering a cube bounded by 6°N, 14°N, 85°E and 92°E (enclosing most of the deep Bay) extending up to 1000m,
the net transports showed an influx of 16.9×1012m3 per month and 124.8×1012m3 per month during the summer and winter monsoons, respectively (Rao & Murty
1992). But this estimate can be considerably different from the real values since the area between 80°E and 85°E, and 5°N and 11°N is one of strong and deep
currents (Figs 42, 43) during both summer and winter. Murty et al. (1993) computed geostrophic currents and transports across 12°N during April and found a
southward transport of 7–8×1012m3 per month for the upper 1000m layer.
Summary
The physical oceanography of the Bay of Bengal and Andaman Sea is reviewed. The review is carried out by dividing the area into two regions: (a) coastal Bay up to
250–300km and (b) deep Bay with the remaining area up to 5°N influenced by river runoff in the coastal areas. The past research activities in the Bay can be grouped
into three periods: (1) those carried out prior to IIOE (2) those carried out during and after IIOE and (3) those carried out after commissioning the first research vessel
of India (RV GAVESHANI, 1976). Most of the studies before IIOE were limited to the coastal areas of the central east coast of India; most of the open ocean data
were collected during and after IIOE.
The Andaman Sea and its coastal areas are studied representatively during the winter monsoon only. Similarly, the coasts of Bangladesh and Burma (northeastern
Bay) are poorly sampled. Data coverages for the Bay of Bengal during pre and postsummer monsoon periods are also poor. The areas north of 13°N in the Bay of
Bengal have not been sampled during the postsummer monsoon period.
One of the important characteristics of the coastal Bay of Bengal and Andaman Sea is dilution of the surface water (extending also into the deep Bay) by the huge
seasonal river runoff from the three river systems, viz., (a) KrishnaGodavari on the central east coast, (b) MahanadiGangesBrahmaputra at the head of the Bay, and
(c) IrrawadySalween on the central east coast of the Bay. Considering the full surface Bay up to 5°N, there is a negative water balance (−11.5cm/season) during
winter and a positive balance (+63.7cm/season)
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during summer. Hence the surface hydrography, especially in the coastal Bay, is greatly influenced by the two monsoons. The 33.0ppt isohaline that runs parallel to the
Indian coast, fully during winter and partly during summer, clearly demarcates the shelf and deep Bay region. Strong winddriven up welling and sinking are seen along
the southern Indian coast and moderate downwelling along the northwestern Indian coast.
Property distributions at the surface show a gradual gradient from the head of the Bay to the outer Bay; from 29.5°C (25.5°C) to 28.0°C (28.5°C) in the case of
temperature and from 21. 0ppt (32.2ppt) to 34.5ppt (34.0ppt) for salinity during the summer monsoon (winter monsoon). During presummer monsoon, the thermal
gradient is rather high (3°C) from the head of the Bay (27.5°C) to the central Bay (30.5°C) and the salinity gradient is small (1.4ppt) from 32.4 to 33.8 ppt. The
southwestern Bay is characterized by high salinity surface waters in all the seasons under the influence of equatorial waters flowing into the Bay. Generally, the surface
salinity distributions are found to follow the current patterns. High salinity water masses flow into the Bay of Bengal at subsurface (50–100m) and intermediate (200–
900m) depths and the latter forms the high salinity (35.0 to 35.1 ppt) core layer of the Bay of Bengal. The thickness and distribution of this layer, however, show wide
seasonal variations. Large horizontal gradients in temperature are noticeable over 100m surface in all the seasons and in turn influence the surface layer circulation.
Horizontal gradients in salinity at 100m are more or less the same as at the sea surface within seasons but drastically decrease from that at the sea surface from one
season to another. At deeper levels, from 200 m, temperature variations are nearly uniform for all the seasons although the locations of the warmwater cells change
with the season. Presence of a largescale subsurface (100m) divergence in the central Bay of Bengal along the axis (southwestnortheast) of maximum surface winds is
an interesting feature during summer and winter with shallowing and deepening, respectively, of the mixed layer.
Data and information on surface and subsurface currents in the Bay and Andaman Sea are very scattered. The situation in deeper layers is still not well known;
practically no studies have been done on bottom currents. Coherent, mean seasonal patterns for the surface currents for the two main seasons (summer and winter) are
derived from the published results. The circulations in the Bay at surface, 500m and 1000m are constituted of steady seasonal gyres, transient cells and eddies. A very
important aspect of the circulation at 500m in the northwestern Bay is its reversal during both seasons. No model would be able to simulate the circulation correctly
unless the freshwater inflows from the three river systems and the currents in the deep channels along the AndamanNicobar chain of islands are suitably incorporated.
Considering surface winds alone (for the area >5°N) MarchApril and OctoberNovember are transition months during which the steady wind pattern of one season
reverses to another pattern. The peak development of the surface circulation in the northwestern Bay seems to occur late during FebruaryMarch and not during the
peak winter period of DecemberFebruary. This can be an effect of the remote forcing from the equatorial area.
The whole scenario of water structure in the Bay involving changes in variations of insolation, precipitation, wind, runoff, evaporation, temperature, salinity, currents,
vertical motions, mixing etc. can be summarized as follows. The water column from surface to bottom can be divided into three layers: (a) surface layer (up to c.100m)
(b) thermocline layer (up to c.1000m including seasonal and oceanic thermoclines), and (c) deep layer (>1000m). The surface layer would absorb almost all the surface
energy inputs and seasonal variations of the mixed layer. The thermocline almost coincides with the intermediate high salinity layer wherein large variations of flow of
high saline waters from the equatorial Indian Ocean
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are found. This layer also accounts for large fields of convergences and divergences. The deep layer is uniform with standard oceanic features with the northward flow
of deep water and its subsequent ascent into the top thermocline (around 400m)
Acknowledgements
The authors thank Dr E.Desa, Director, National Institute of Oceanography for supporting this work. We record our thanks to all the authors and publishers who
agreed to reproduction of some figures from their works. We would like to state here that omissions of past works, if any, are unintentional.
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NUTRIENT FLUXES TO PLANKTONIC OSMOTROPHS IN THE

PRESENCE OF FLUID MOTION
L.KARPBOSS, E.BOSS & P.A.JUMARS
University of Washington, School of Oceanography, Box 357940, Seattle, WA 98195–7940, USA

Abstract We present solutions for nutrient transfer to osmotrophs in the full range of flow regimes for which solutions have been published, and we
extend some of those solutions to new parameter domains and flow environments. These regimes include stagnant water; steady, uniform flow arising
from swimming or sinking; steady shear flows; and fluctuating shear from dissipation of turbulence, as well as the combined effects of turbulence
induced shear and swimming or sinking. Solutions for nutrient fluxes cannot be carried over from one flow regime to another. In all cases, however,
mass transfer increases with cell size and with flow velocity. Cell shape becomes particularly important at high flow velocities. For steady, uniform flow
arising from sinking or swimming, we find asymptotic analytic and numerical solutions from the engineering literature superior to those in more common
use within oceanography. These engineering solutions suggest flow effects an order of magnitude smaller than commonly supposed. A cell radius near
20μm is needed before swimming or sinking can be expected to increase the flux of nutrients, such as nitrate or phosphate, substantially (by ≥50%)
over the stagnantwater case. We find sound asymptotic solutions for the case of linear shear and supplement them with numerical solutions of our
own to cover the range of cell sizes and shear rates of interest for phytoplankton. We extend them further to cover viscous shears from dissipating
turbulence for cells smaller than the Kolmogorov scale (order of 1–6mm in the ocean). Our analysis suggests turbulence effects an order of magnitude
greater than previously postulated, with a cell size of 60μm needed to experience substantial gain. Cell rotation, whether induced by the propulsion
mechanism in swimming or passively by shear across the cell perimeter, will reduce the rate of nutrient transfer relative to a nonrotating cell unless the
axis of rotation parallels the direction of flow. Although in calm water dinoflagellates by swimming are able to increase nutrient uptake, in strong
turbulence they may not be able to maintain a rotational axis parallel to the direction of swimming or the direction of shear, resulting in a relative
reduction in flux. Conversely, large chains of diatoms and filamentous cyanobacteria that span the radius of the smallest vortices are best able to take
advantage of turbulence. Despite these deductions from a diversity of analytic and numerical solutions, unequivocal data to test the contribution of
advection to nutrient acquisition by phytoplankton are scarce—owing, in large part, to the inability to visualize, record and thus mimic fluid motions in
the vicinities of cells in natural flows.
Introduction
Effects of smallscale fluid motions on diffusional fluxes of nutrients to bacteria and phytoplankton have been reviewed several times (Gavis 1976, Roberts 1981, Lazier
& Mann 1989, Mann & Lazier 1991, Kiørboe 1993). It is thus worth asking why it might be worthwhile to visit them yet again. In the process of a review focused
much more closely on bacteria, we (Jumars et al. 1993) became aware of several inconsistencies in the published treatments of solute fluxes to osmotrophs. Among the
accumulated problems we encountered, one solution (Munk & Riley 1952) is often used for sinking cells while another (Berg & Purcell 1977) is often used for
swimming cells, despite the fact that both invoke the identical governing equation and boundary conditions and thus can neither apply to disparate
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processes nor both be correct. Unfortunately, we perpetuated several other inconsistencies. In particular, we were not adequately aware of the diversity of field and
laboratory flow regimes for which solutions had been given or of the fact that competing solutions had been published for some regimes. The principal purpose of our
present review, therefore, is to lay out the diversity of flow regimes plainly and to point out the best of the known solutions for each specific case. We focus on those
physical characteristics of the cell and its flow environment that affect nutrient flux.
A secondary purpose is to specify, as accurately as can be done from the existing literature, the size of organism for which relative fluid motion begins to become
important in nutrient flux. It has been firmly established that, for diffusion coefficients typical of small molecules in water (of order 10−5cm2s−1), 1μm long, freely
suspended bacteria cannot experience any substantial flux enhancement from the fluid shear about them (Berg & Purcell 1977, Lazier & Mann 1989, Logan & Dettmer
1989). Utilization of slowly diffusing molecules, larger “cell” size or attachment to larger particles, however, can result in significant benefit from relative fluid motion
(Roberts 1981, Logan & Hunt 1987, 1988, Lazier & Mann 1989, Logan & Dettmer 1989, Confer & Logan 1991). The ranges of sizes and diffusion coefficients for
which such enhancements can become important have been given implicitly or explicitly in several treatments (Logan & Hunt 1987, 1988, Lazier & Mann 1989), but
have sometimes been based on equations from inappropriate flow regimes. As with many other realworld biologicalphysical interactions of interest, these substantial
enhancements of fluxes begin in an uncomfortable middle range of parameters for which analytic solutions typically are not available. Our systematic exploration
identifies flow regimes and parameter values for which further numerical modelling and laboratory experimentation would be profitable.
The role of fluid motion in the nutrition of planktonic autotrophs and osmotrophs is similar to its role in problems of heat and mass transfer long studied by civil and
chemical engineers. Since the governing equations and boundary conditions for the heattransfer problem and the dilutesolution masstransfer problem are identical,
heattransfer theory is applicable directly to the study of nutrient fluxes to planktonic osmotrophs. Most of the solutions we present thus come from the engineering
literature; we use, however, oceanographic (Appendix I) rather than engineering convention for notation. In keeping with practice in physics and engineering, we use
square brackets to denote primary dimensions of length [L], mass [M] and time [T]. To avoid confusion, we avoid the term “convection” and label any organized fluid
motion as “advection”. In this review diffusion and molecular diffusion are synonymous. Several inconsistencies have occurred because some authors used radii while
others used diameters as characteristic dimensions; for consistency, we give characteristic cell sizes as radii. We work from stagnant fluids, through simple, steady shear
flows to the threedimensional, rotational shear flows characteristic of dissipating turbulence. We warn fluid dynamicists that we use the word “particle” exlusively for a
solid object of finite dimensions suspended in the flow and not synonymously with a small parcel of water.
We focus on phytoplankton, because it spans the full size range of interest (from a few micrometres to a few millimetres) and comprises a wide diversity of body,
chain and colony shapes. Our analysis also applies, however, to metazoan larvae and other freeliving planktonic osmotrophs. To save space we refer to the individual
as a cell and leave implicit the applicability to multicellular organisms of comparable sizes. We also leave implicit the converse problem of solute leakage from a cell,
although it is plain (Jumars et al. 1993, their Fig. 1) that our results apply equally well there. The principal difference is a sign change in the concentration gradient and
hence in the flux direction; magnitudes of flow effects on such
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fluxes are identical. We also caution that solutions frequently given for a point source or sink cannot adequately describe the problem of solute leakage from or uptake
by a cell; an important aspect of fluid motion is the divergence of streamlines around cells.
Mass transfer to cells in stagnant water
Nonmotile cells: pure diffusion

Rate of nutrient delivery into a planktonic cell at steady state proceeds at the rate of the slower of two processes in the series, nutrient delivery to the cell membrane
and transport across the membrane. If the former is much faster, no nutrientdepleted layer develops around the cell. If the latter is faster, however, nutrient
concentration at the cell surface drops below its farfield level, thereby driving a faster delivery rate by diffusion.
The concentration distribution of a nutrient about a planktonic cell in either case is given by the diffusion equation (Crank 1975),
(Equ. 1)
where D is the diffusion coefficient [L2 T−1] and C is the concentration of the nutrient of interest. The del operator ( ) indicates partial derivatives in all three spatial
dimensions (e.g. in Cartesian coordinates ). In a heterogeneous environment like the ocean, D varies in time and space (due to temperature and
salinity). Since we are interested in processes at the microscale, D for a given nutrient is assumed to be constant. Thus, (D C)=D 2C.
At steady state the solution of Equation 1 for a sphere becomes:
(Equ. 2)
where r0 is the cell radius, r is the radial distance from the centre of the cell, C∞ is the farfield concentration and C0 is concentration at the cell surface. When uptake
capacity into the cell exceeds diffusional supply rate, Equation 2 indicates that there exists a nutrientdepleted region in the vicinity of the cell (Koch 1971, Jumars
1993). For convenience, hereafter we refer to this depleted region as the diffusional boundary layer and define it more specifically as the region in which C≤90% of the
ambient concentration. By this definition it extends to about nine cell radii away from the cell surface (Fig. 1 herein; Koch 1971, Jumars 1993). The outer limit is
arbitrary, since approach to background concentration is asymptotic. The rate at which nutrients will be transported across this diffusional boundary layer can be
estimated from the characteristic diffusion timescale (tD),
(Equ. 3)
where Lc is a characteristic length scale (e.g. the thickness of the layer). For a cell with a radius of 10μm and a diffusional shell 90 μm thick, it will take c.8s for a
molecule such as nitrate or phosphate (D of the order 10−5cm2s−1) to cross this layer.
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Figure 1 Concentration distributions of a given nutrient around cells (black circles) in three different flow regimes: stagnant water (pure
diffusion), uniform flow (i.e. cell swimming or sinking in stagnant water; the cell is moving from right to left so flow is from left
to right) and shear flow (uniaxial extensional flow where E=(2, 0, 0; 0, −1, 0; 0, 0, −1). The third dimension can be visualized by
rotation about the x axis. The left panel illustrates the concentration field at distances up to 10 cell radii from the centre of the
cell. The right panel is a “blowup” of the concentration field near the cell surface, up to a distance of 2 radii from the centre of
the cell. Concentration values for the case of pure diffusion were obtained by solving Equation 1 at steady state analytically.
Values for the two other cases were obtained by solving Equation 11 numerically for Pe=1 to steady state (Appendix II). In the
absence of fluid motion a diffusive boundary layer (C≤0.9C∞ ) extends to about 9 cell radii from the cell surface. Uniform and
shear flows distort the boundary layer and steepen the concentration gradient in certain regions. Since transport is dominated by
the thinnest regions of the diffusional boundary layer, cells in uniform or shear flow will experience enhanced fluxes compared
to nonmotile cells in stagnant water.
The diffusional flux of nutrients to a spherical cell (QD) depends on the diffusion coefficient, th e cell radius (r0) and the concentration gradient between the cell surface
and the ambient water,
(Equ. 4)
There are only two means by which the cell can increase the flux to itself without relative fluid motion: increasing r0 and lowering C0 (Jumars et al. 1993). There are,
however, limits
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on the increase achievable. As cell size increases, demand for nutrients increases more rapidly (proportional approximately to r02 cf. Roberts 1981) than does the
diffusive flux. In addition, for a given cell size, it is impossible to enhance the flux beyond its value for C0=0, when every molecule that reaches the cell surface is
absorbed immediately.
Assuming delivery to a spherical cell by molecular diffusion and active uptake of MichaelisMenten form across the cell membrane, Pasciak & Gavis (1974)
introduced a criterion for recognizing transport limitation. They defined the parameter P to be the ratio between diffusional delivery and maximal uptake rates,
(Equ. 5)
where Vmax is the maximal uptake rate [mol cell−1 T−1] and Km is the concentration at which the uptake rate equals ½Vmax. When P is small, the cell has the capacity
to absorb nutrients much faster than the rate at which nutrients can diffuse towards the cell, and transport to the cell membrane becomes limiting. When P is large, the
maximal uptake rate is much slower than diffusion, and for such an organism uptake rate will not be controlled by its physical environment. Pasciak & Gavis (1974)
applied this criterion to phytoplankton, for which values of Km and Vmax were known from laboratory experiments, and thereby established that for some
phytoplankton (mostly large cells) transport limitation can be significant. It has also been suggested that under certain conditions uptake of carbon dioxide (Riebesell et
al. 1993) and iron (Morel et al. 1991) by phytoplankton and uptake of phosphate by cyanobacteria (Mierle 1985) may become transport limited.
Determining whether uptake plays a role in limitation is clearly important but is outside the scope of our review. For all subsequent analyses, we assume that the cell
is a perfect absorber, i.e. that cellsurface nutrient concentration (C0) equals zero. We do so for two reasons. One is to provide an upper bound on the importance of
fluid dynamic effects: if there is no significant effect of fluid motion on uptake for a perfect absorber, then there can be none for an imperfect one. The more obvious
reason is for compactness. If kinetics of uptake are known and suspected to be important, they can be included in the problem in the manner shown by Pasciak &
Gavis (1974, 1975).
Motile cells: swimming and sinking
Theory
Swimming and sinking are two mechanisms by which a phytoplankter can induce relative fluid motion, steepen its surrounding solute gradients and hence increase the
fluxes of nutrients to itself. The nature of the flow in the vicinity of the cell is determined by its body Reynolds number (Re). This Reynolds number describes the relative
importance of inertial forces compared to viscous forces,
(Equ. 6)
where U is the characteristic velocity, v is the kinematic viscosity [L2 T−1] and Lc is the characteristic length scale, in our case the cell radius (r0). Since typical values of
Re for plank
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Table 1 Reynolds and Péclet numbers for swimming and sinking phytoplankton.
Organism Cell radius Swimming speed Sinking speed Re Pe Source

(μm) (μms−1) (μms−1)
Prorocentrum marial 6 171 0 1×10−3 1 Kamykowski et al.
labouriae (1992)
Gyrodinium dorsum 16.5 330 0 5.4×10−3 5.4 Kamykowski et al.
(1992)
Dinophysis acuta 33 500 0 1.6×10−2 16.5 Sommer (1988)
Coccolithus huxleyi 5 to 6.5 0 3 to 15 1.5×10−5 to 1.5×10−2 to Smayda (1970)
9.7×10−5 9.7×10−2
Coscinodiscus wailesii 70 0 81 to 347 5.7×10−3 to 5.7 to 24 Smayda (1970)
2.4×10−2
tonic microorganisms are well below unity (Table 1), their environments are dominated by viscous forces, and inertial forces can be neglected.
In the presence of fluid motion the concentration distribution of the nutrient of interest around an organism is given by
(Equ. 7)
where U is the threedimensional velocity field. As a first approximation, we use Stokes’ solution for uniform, creeping flow past a stationary, rigid sphere (Leal 1992,
Ch. 4) to describe the flow field around cells swimming or sinking in stagnant water. To permit analytic determination of the velocity field and concentration distribution
in the neighbourhood of the cell, we assume spherical shape and sufficient spacing between cells to preclude interactions among them.
The boundary conditions applied to Equation 7 are:
(Equ. 8)
(Equ. 9)
These boundary conditions set an upper bound for the calculated flux. The flux of a given nutrient (Q) to the cell is determined by the integrated concentration gradient
normal to the cell surface:
(Equ. 10)
where A is the cell surface and n is an inward unit vector normal to it.
Much of the information and insight about key parameters that determine the concentration distribution in the presence of fluid motion can be obtained via
nondimensionalization, without the need to solve the differential equation. Defining U*=U/U; C*=C/C∞ and X* , Y* , Z* =x/r0 , y/r0 , z/r0, the nondimensional form of
Equation 7 at steady state becomes
(Equ. 11)
where each asterisk denotes a nondimensional variable. Corresponding boundary conditions are:
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(Equ. 12)
(Equ. 13)
for r* the nondimensional radial distance from the centre of the cell (r* = r/r0) and Pe, the dimensionless Péclet number. The Péclet number indicates the effectiveness
of advective transport compared with diffusive transport through the fluid over the specified length scale:
(Equ. 14)
where U is the characteristic velocity (swimming and sinking velocities in the case of motile cells). Diffusion dominates transport when Pe<1, and advection dominates
when Pe>1. Note that while Reynolds numbers for swimming and sinking planktonic microorganisms are always smaller than unity, Péclet numbers for large cells can
be larger (Table 1). Again, the body Reynolds number of an organism will determine the flow field in its vicinity while the Péclet number will determine the concentration
field of nutrients and hence the fluxes.
Since Pe is the only nondimensional parameter arising from the steadystate form of Equation 7, any desired nondimensional quantity determined by the
concentration field, namely the nondimensional flux, will be dependent solely on Pe. A measure of this flux is the Sherwood number, Sh (its heattransfer analogue being
the Nusselt number, Nu), which is the ratio between the total flux of nutrients arriving to the cell surface in the presence of fluid motion (Q) and the purely diffusional flux
(QD):
(Equ. 15)
Sh thus indicates relative enhancement of flux due to advection. In the case of a spherical cell in stagnant water Q=QD and Sh=Sh0 =1
In the engineering literature the Sherwood number is commonly defined alternatively in terms of the masstransfer coefficient, k, [L T−1] as Sh 2kr0 /D. In this case,
Sh0 =2. The masstransfer coefficient is the ratio between the mass flux per unit of area and the concentration gradient between the boundary of interest and the
environment (Cussler 1984, Rohsenow & Choi 1961). It can be determined empirically and varies with flow regime and shape of the body. This definition has been
adopted in oceanographic applications but in the form Sh kr0 /D (Sh0 =1, e.g. Logan & Hunt 1987, 1988). The difference arises from selection of the diameter versus
the radius as the characteristic length scale. For the same reason, and hence by the same factor, definition of the Péclet number in the engineering literature often differs
from the oceanographic convention, i.e. in the engineering literature it is often Pe 2Ur0 /D. To reduce confusion among oceanographic readers, we have converted all
formulas extracted from the engineering literature to the oceanographic convention of Equations 14 and 15 (Appendix I). We make this point and our parameterizations
explicit because factors of two based on choice of diameter versus radius as the characteristic dimension have caused some mischief in past comparisons.
In order to examine the effect of fluid motion on nutrient flux a functional relationship between Sh and Pe is sought. First it is necessary to find a solution to Equation
11 that satisfies the boundary conditions 12 and 13. Unfortunately, this problem is too complex to permit exact solution over the full range of Pe. One way to approach
it analytically is to derive
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asymptotic solutions for limiting values of Péclet numbers. Asymptotic relations between Sh and Pe have been derived for the two extreme cases, Pe 1, when
advective effects are relatively minor compared to pure diffusion and Pe 1, when the effect of molecular diffusion may be neglected everywhere except for a thin
boundary layer at the fluidcell interface. The major difference between the two is the nature of the dependence of the concentration distribution on flow geometry and
hence the nature of the dependence of Sh on Pe. For low Péclet number, the concentration distribution near the body is governed by molecular diffusion and is
therefore relatively insensitive to flow geometry within that region. Cell surface relief smaller than 0.1r0 in dimension can have no great effect in this regime. At large
distance from the body, however, the form of the velocity field determines the concentration field, so overall body shape and orientation can be influential. At large
Péclet number, on the other hand, the effect of molecular diffusion is limited to a very thin layer around the cell, and therefore the concentration distribution will be very
sensitive to the flow field in the vicinity of the cell (Leal 1992). Here, smallscale relief on the cell surface can have profound effect. For this reason, flux will be much
more sensitive to finescale details of cell shape at high Pe than at low.
Figure 2 Sherwood number (Sh) as a function of Péclet number (Pe) for cells moving at a constant velocity in stagnant water or cells
fixed in a uniform flow (Re 1). Equation 16 was derived by Acrivos & Taylor (1962) for Pe 1 and Equation 17 was derived
for Pe 1 by Acrivos & Goddard (1965). Clift et al. (1978) suggested Equation 18 as a fit to their numerical results; we use it
for the region of intermediate Pe for which analytic solutions are not available. Berg & Purcell (1977) obtained their relation
numerically, but provided no explicit equation. For reasons detailed in the text and in Appendix II, we believe Berg & Purcell’s
(1977) relation (their Fig. 4) to be inaccurate.
Asymptotic solutions for Equation 11 in the case of steady, uniform, laminar flow and the corresponding boundary conditions (12, 13) yield two relations between Sh
and Pe (Fig. 2). For Pe 1 (Acrivos & Taylor 1962)
(Equ. 16)
For Pe 1 (Acrivos & Goddard 1965)

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(Equ. 17)
where O(x) is read as “of order x”, meaning ≥0.5x but ≤5x, the precise coefficient varying with the details of the situation, and o(x) indicates a term much smaller than
the term in brackets. Equations 16 and 17 were derived for the case of heat transfer from a spherical particle and were given in the form of the Nusselt number (Nu).
For detailed considerations of the use of asymptotic techniques in the analysis of heatandmass transfer problems and for lucid derivations of the above solutions we
refer the reader to Leal (1992).
Péclet numbers for many if not most phytoplankton fall outside the region for which analytic solutions are available (Table 1). Numerical solutions that include the
region of our interest were published by Clift et al. (1978) and Masliyah & Epstein (1972). The former suggested the following relationship for Sh, valid for all Pe in
creeping (Re<0.1), uniform flow:
(Equ. 18)
So long as very low Re is maintained, the above formula agrees within 0.7% in the region 0.001≤Pe≤0.1 (Equation 16) and within 2% of the analytic solution (Equation
17) in the region 100≤Pe≤5000.
Application of the theory to swimming and sinking cells

Most classes of marine phytoplankton are motile (Sournia 1982). The most thoroughly studied motile phytoplankters are the dinoflagellates; their swimming speeds
range from 50 to 500μms−1 (Throndsen 1973, Sournia 1982). A nonmotile phytoplankter can sink or rise due to density differences between itself and the
surrounding fluid and thus also experience a velocity relative to its environment. For cells of a given size, measured sinking speeds are usually slower than swimming
speeds and range from 0 to 347μms−1 (Smayda 1970, his Table VII).
To illustrate the effect of swimming or sinking on the concentration distribution of a given nutrient we solved Equation 11 with the corresponding boundary conditions
(12, 13) numerically for Pe=1 (Fig. 1; Appendix II). For both high and low Pe, a disproportionate amount of the total flux takes the path of least resistance. As can be
seen by inspection of Equation 10, nutrient transport will be dominated by the thinnest regions of the diffusional boundary layer, where diffusion can proceed by far the
fastest; characteristic diffusion time varies with the square of this thickness (Equation 3). Hence, total flux arriving to the cell will be larger compared with the purely
diffusional flux despite the thickening of the diffusional boundary layer at the “rear” of the cell. Again, some swimming modes may help to “shed” this thickened
boundary layer or to thin the thinnest regions even further compared with steady, uniform, Stokes’ flow past a sphere. Interestingly enough, it has been reported that
chemoreceptors on the cell surface of the motile bacterium Escherichia coli are clustered at the pole of the cell rather than being evenly or randomly distributed
(Maddock & Shapiro 1993). Such clustering may increase the efficiency of detection, especially if the receptorrich patch is located at the “leading” pole of the cell
(Parkinson & Blair 1993) or where a flagellum locally thins the boundary layer. The advantage is greatly decreased detection time. Berg & Turner (1995) observed that
tumbles tend to randomize cell orientation of E. coli and that runs of each orientation are of similar duration. They therefore concluded that patches of chemoreceptors
at one pole of the
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cell do not serve as the bacterium’s “nose”. Clustering of chemoreceptors or uptake sites at the poles or at other sites of local boundarylayer thinning, however, may
still be an effective strategy for other organisms with different modes of swimming.
In order to quantify the enhancement of flux due to swimming and sinking, for the full range of Pe values and cell sizes, we use the solutions for mass transfer in
steady, uniform flow—namely Equations 16–18. The only differences in our proposed solutions for sinking versus swimming (Fig. 3) arise from differences in the
dependence of settling versus swimming speeds on cell size and hence the dependence of Pe and Sh on cell size. We base swimming speeds on the relationship,
(Equ. 19)
Figure 3 Sherwood number (Sh) as a function of cell radius (r0) for swimming and sinking cells. Both swimming and sinking velocities
are functions of cell size; Stokes’ equation was used to calculate sinking velocities and the relation U=αr0β was used to
calculate swimming velocities. We caution that some correlations between cell size and swimming velocity are very weak; we
use them only for illustration. Dashed lines represent sinking cells and solid lines represent swimming cells where: a, sinking
cells Δρ=0.01; b, swimming cells U=0.16(2r0)0.46 (Kamykowski et al. 1992, R2=0.32, based on 5 species, ranging in size (r0)
from c. 13 to 30μm); c, swimming cells U=93(2r0)0.26 (Sommer 1988, r=0.38, based on 19 species, ranging in size (r0) from c.
1 to 40μm); d, sinking cells Δρ=0.1; and, e, swimming cells U=30.66(2r0)1.16 (Kamykowski et al. 1992, R2=0.79, based on 9
species, ranging in size (r0) from c. 8 to 25μm). To calculate Sh we used equation 16 for Pe<0.01, Equation 18 for 0.01
<Pe<100 and Equation 17 for Pe>100. The diffusion coefficient for the calculations of Pe was taken to be 1×10−5cm−2s−1.
The gray segments of curves a, d, and e mark regions for which swimming and sinking velocities extrapolated from the given
relations become unrealistic.
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which was obtained by Sommer (1988) from the empirical data of Throndsen (1973) and Sournia (1982) and by Kamykowski et al. (1992) from the empirical data of
Kamykowski & McCollum (1986). Coefficients α and β vary among species, where 0<β<2. We use this function as an example but are aware that for some species
its fit is poor. For sinking velocities we used Stokes’ equation for sinking spheres (not to be confused with Stokes’ solution for the flow field),
(Equ. 20)
where g is the gravitational acceleration constant (980cms−2), ρc and p are the densities of the cell and fluid, respectively [ML−3], and μ is dynamic viscosity
(approximately 0.01gcm−1s−1; for more accurate values as a function of temperature and salinity see Jumars et al. 1993). Excess densities (ρc−ρ) used for the
calculations were 0.1 and 0.01 gcm−3 (Van Ierland & Peperzak 1984). The diffusion coefficient (D) for nutrients such as nitrate and phosphate is O(10−5)cm2s−1. For
calculations of Péclet number, D was taken to be 1×10−5 cm2s−1.
For both swimming and sinking, nondimensional mass transfer to the cell depends directly (but not necessarily linearly) on cell size and cell velocity relative to the
water and inversely on the diffusion coefficient, as implied from the relationship between Sh and Pe. Realistic swimming or sinking speeds do not increase substantially
the flux of nutrients such as nitrate or phosphate to small cells (Fig. 3). Critical cell sizes for which swimming and sinking begin to become important (to be conservative,
we consider a substantial enhancement as an increase of 50% or more of the flux) will vary with the parameters determining the specific dependence (Table 2).
Table 2 Predicted cell radii for 50 and 100% increase in flux, based on equations 16–18.
Predicted cell radius (mm)

Mode of motility 50% increase in flux 100% increase influx
Swimming, U=30.66(2r0) 1.16 12 22
Swimming, U=93(2r0)0.26 15 44
Swimming, U=0.16(2r0)0.46 25 61
Sinking, Δρ=0.1 25 39
Sinking Δρ=0.01 54 84
Turbulenceinduced shear 63–100 (calculated from the lower and upper limits for 167–202 (calculated from the lower and upper limits for
(ε=10−2cm2sec−3) interpolation; (see Fig. 6) interpolation; (see Fig. 6)
Motility, however, may become beneficial for small cells if larger molecules are considered. Amon & Benner (1994) have shown that highmolecularweight, dissolved,
organic matter (HMW DOM; molecules smaller than 0.1 μm but larger than 0.001μm) is more readily remineralized by bacteria than lowmolecular weight DOM
(molecules smaller than 0.001 μm). Diffusion coefficients (at 20°C) for HMW DOM can range from c. 10−6cm2s−1 to c. 10−8cm2s−1 (estimated from the size cutoff
for HMW DOM following Cussler 1984). For a bacterium 0.45μm in radius, swimming at 20μms−1 will not appreciably enhance the flux of molecules having a
diffusion coefficient O(10−5)cm2s−1 but will increase the flux of
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molecules with a diffusion coefficient O(10−8)cm2s−1 by 80%. The predicted contribution of advective flux also depends on ambient temperature through the latter’s
effect on the diffusion coefficient of the solute (Cussler 1984) and on viscosity (Korson et al. 1969) and thus on sinking velocities and velocity gradients in the vicinity of
the cell. While mixotrophy and particle feeding are outside the scope of our review, we note that the diffusion coefficients used to characterize Brownian motion of
particles are far smaller than diffusion coefficients for molecules, so that fluid motion may have substantial effect (Shimeta 1993, Shimeta et al. 1995).
The theory we have applied (i.e. Equations 16–18) is at odds with previous, widely cited theory that examined the utility of swimming and sinking in enhancing
nutrient fluxes. Munk & Riley (1952), who examined effects of sinking on nutrient absorption by phytoplankton cells of varying shapes, applied an analytic solution
derived by Kronig & Bruijsten (1951) for spheres in uniform flow at Pe 1:
(Equ. 21)
Other solutions were not available to them at the time. For the firstorder approximation, the correlation they used agrees with Equation 16. For larger values of Pe,
Munk & Riley (1952) used the empirical relationship given by Kramer (1946), which is valid only for Re>1 and therefore not for sinking phytoplankton. No explicit
relation between Sh and Pe was given by Munk & Riley (1952) for the region of intermediate Péclet numbers. Their estimates for that region were obtained by
interpolation between the analytical and empirical relationships.
Berg & Purcell (1977), who examined the effect of swimming on nutrient uptake, solved Equation 11 with boundary conditions 12 and 13 numerically. They did not
provide an equation for the relation between relative increase in mass transfer and Péclet number, but their model results suggested that for Pe 1 the increase in
nutrient flux is proportional to Pe2, while for the case of Pe>1 (though they did not examine the increase of flux beyond Pe=10) the flux is proportional to Pe1/3. Berg
& Purcell (1977) argued that the increase of flux in the neighbourhood of U=0 must depend on an even power of U since it cannot depend on the direction of the
motion and cannot have a singularity at U=0. The total nondimensional flux, integrated over the whole surface area of a spherical cell, however, does not depend on
the direction of the flow—only on the magnitude of the velocity (as implied from the relation Sh =f(Pe)), and therefore we find no particular reason to argue for
dependence of an even power. Moreover, the shape of the curve implied by Berg & Purcell’s correlations (the exponent of Pe going from 2 to 1/3, i.e. a sigmoidal
curve) implies an intermediate Péclet number for which the fractional increase of flux (δSh/δPe) is maximal. Since we are dealing exclusively with small Re, changes in
the flux as a function of Pe are expected to be gradual and monotonic. We cannot find any physical basis for steeper change at any particular velocity, cell size or
diffusion coefficient. We speculate on the source of the discrepancy between Berg & Purcell’s (1977) numerical solution and the analytical solution or our numerical
solution in Appendix II.
Berg & Purcell’s estimate for the increase in the nondimensional flux due to advection is higher than predicted by Equation 18 in the region Pe>0.04 (Fig. 2). For
example, Berg & Purcell estimated c. 76% increase in the flux for Pe=1 while Equation 18 and the results of our numerical model (Fig. 1; Appendix II) predict an
increase of 22 and 21 %, respectively. In order to double the flux of nutrients to the cell, the Péclet number should be of the order 2.5 according to Berg & Purcell
(1977) and 13 according to Equation 18 (Fig. 2). Estimates of the effect of swimming on nutrient transfer (Roberts 1981, Goldman 1984, Sommer 1988, Lazier &
Mann 1989, Mann & Lazier 1992) that were calculated from the graphical presen
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tation of Berg & Purcell (1977) must, therefore, overestimate the contribution of motility to nutrient fluxes. Rather than endangering Berg & Purcell’s central conclusion
that swimming will be ineffective in enhancing the flux of rapidly diffusing (D=10−5cm2s−1) nutrients to bacteria, the analytic solution makes it even more unassailable.
Our results (Fig. 3) indicate that there is no one mechanism that is always more beneficial in enhancing the flux, and call into question Gavis’s (1976) conclusion that
swimming is more effective than sinking in reducing diffusional transport limitation. Other factors, such as the cost of swimming and the risk of being transported away
from the euphotic zone when sinking rapidly, should be taken into account, however, when the benefits of each mode are considered. Dependence of swimming speed
on cell size varies with mode of self propulsion, i.e. planar wave motion, helical wave motion or a combination of both. For instance, cells driven by a flagellum beating
with a helical wave rotate as they swim (Chwang & Wu 1971, Brennen & Winet 1977). Smaller cells tend to have higher rotational velocities that slow their
translational velocities compared to larger cells. As cell size increases, however, drag increases, too, and larger cells that may escape the problem of rotation face
increasing drag, which tends to reduce their translational velocities. Thus, for a phytoplankter using this mode of swimming there is an intermediate size for which
swimming speed is maximal (Brennen & Winet 1977, Kamykowski & McCollum 1986). Sinking speeds may also deviate from the predicted relation with cell size.
Empirical data on sinking speeds as a function of cell size indicate that the slope can be smaller than the value of 2 predicted by Stokes’ law, no doubt the result of a
correlation between size and excess density (Waite et al. 1992). Sinking or rising speeds may vary with nutrient and light conditions (Culver & Smith 1989) and depend
on the growth phase of the cell (Eppley et al. 1967, Smayda 1970). Among the majority of planktonic diatoms, nutrientdepleted cells sink more rapidly than nutrient
replete cells (Titman & Kilham 1976, Smetacek 1985, Waite et al. 1992). Within a species there may be variations in sinking speeds of a factor of two or more with
varying nutritional status of the cell, accompanied by relatively small changes in cell size (Jackson & Lochmann 1993). We have applied those correlations,
nevertheless, to demonstrate that effects of sinking and especially swimming on the flux of nutrients cannot be generalized and that the dependence of Sh on cell size will
vary with the mode of motion (Table 3).
Effects of selfinduced rotation vary with orientation of the rotational axis and with Pe. Rotation, if strong, can reduce advective contribution to flux when it causes
streamlines to curve and sometimes even to close. Water can circulate around the cell longer before being replaced by new water. Advective flux to a rotating,
swimming cell will decrease relative to a nonrotating swimmer, however, only if the cell has a component of translation in a direction perpendicular to the axis of
rotation, and the decrease will be most important at high Pe when advection dominates transport. At low Pe diffusion is the dominant transport mechanism, and rotation
will distort the diffusive boundary layer around the cell by shearing water parcels in its vicinity (as a result of the noslip boundary condition at the cell surface).
Table 3 Dependence of Sh on cell radius for swimming and sinking cells and for neutrally buoyant cells suspended in turbulent water.
Flow generated by Pe<<1 Pe>>1

Swimming Sh r0 1+β
Sh r0(1+β)/3
where 0<β<2 where 0<β<2
Sinking Sh r0 3 Sh r0
Turbulence/shear Sh r0 Sh r02/3
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This distortion of the diffusive layer can increase diffusional flux relative to a swimming, nonrotating cell. The effect, however, will probably be very small. The axis of
rotation of dinoflagellates propagating in a helical wave is parallel to the direction of motion. By our arguments such a swimming mode will still provide a flux advantage
across the full Pe range, and it appears to carry little cost for most ciliary and flagellar mechanisms (Purcell 1977, Raven 1982, Raven & Richardson 1984, Fenchel
1987) unless very high speeds (in terms of body lengths per unit of time) are achieved (Mitchell 1991).
Our and prior applications of the same flow field (Stokes’ flow) to both swimming and sinking are, at best, rough approximations. Swimming and sinking cells differ
in the nearcell details of flow streamlines around them. While streamlines around sinking cells closely resemble Stokes’ flow, in the near field, streamlines around
swimming cells may differ (Keller & Wu 1977) and will vary among species, depending on the mode of swimming. Precise determinations of flow fields around
swimming cells require numerical models and observations specific to each case. Our application of Stokes’ flow is clearly most questionable for swimming at high Pe.
Motions near swimming appendages will be unsteady, thinning the diffusional boundary locally. Because the diffusion time scale varies with the square of diffusion
distance, such thinning is disproportionately important relative to unsteady thickening in other regions. Hence at high Pe we will underestimate the true Sh and advective
contribution to the flux. Since the details depend on local thinning, there can be no general solution; specific solutions will be time dependent and will also be highly
dependent on the mode of swimming and the shape of the cell. In short, flux to swimming cells appears to be a fertile ground for detailed numerical simulation.
Effect of shape on the nutrient flux

Among phytoplankton, relatively few species are spherical. Morphological diversity in nature suggests that each form has a slight advantage in a particular set of
conditions (Munk & Reilly 1956, Sournia 1982). Grazing, drag and nutrient absorption are the commonly suggested selective factors leading to the observed diversity
(Sournia 1982). The nondimensional flux to a body of an arbitrary shape (i.e. its Sh) is defined as the total flux arriving to the cell surface in the presence of fluid
motion normalized to the purely diffusional flux arriving to a sphere with an equivalent surface area (Leal 1992). A general solution for mass transfer to a body of
arbitrary shape in steady flow for Pe 1 was given by Brenner (1963):
(Equ. 22)
where f is the dimensionless drag on the body [f=(drag force)/6πμr0U∞ , with f=1 for a sphere] and Sh0 is the mass transfer to the shape of interest in the case of pure
diffusion and depends upon the geometry of the body. Sh0 can be estimated for each shape from theory (Clift et al. 1978, their table 4.2) or by experiment.
Dimensionless drag, f, depends on orientation of the body relative to its direction of motion. From experience at high Re, one might assume that a sinking cell would
orient itself such that the drag is maximized (for example, that a cylindrical cell would sink with its largest projected area perpendicular to the falling axis). For Re<0.1,
however, the initial (arbitrary) orientation will be kept during sinking (McNown & Malaika 1950) unless weight is unevenly distributed in the cell (Hutchinson 1967,
Sournia 1982). Values and expressions
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for the resistance of spheroids in uniform flow were given by Happel & Brenner (1965, their table 4–26.1) and Clift et al. (1978, their table 4.1).
For Pe>>1 the relationship Sh=cPe1/3 holds for rigid bodies of arbitrary shapes where the coefficient c varies with the geometry of the body but is always of order
unity (Friedlander 1957, Clift et al. 1978, Leal 1992). For the intermediate region of Péclet numbers, numerical results for spheroids were published by Masliyah &
Epstein (1972). These correlations between Sh and Pe are only valid, however, for simple, smooth shapes without spines, horns and sharp corners (Leal 1992). Spines
and horns are common among phytoplankton and may have important roles in altering flow fields around cells and thus fluxes to cells. It is commonly thought that eddy
formation does not occur at Re<1, but eddy formation has been demonstrated within a corner formed by two intersecting planes in creeping flow (Jeffrey & Sherwood
1980). Eddies will reduce the contribution of advective transport since they will cause water to remain longer within that region before being replaced. These studies
may have relevance to some diatoms such as Thalassionema and Asterionella species that form starshaped colonies or zigzag bands of colonies. Also pertinent to
sinking chains is the study by Dorrepaal & O’Neill (1979) for the case of uniform Stokes’ flow past two parallel, separated cylinders with flow direction perpendicular
to the line joining their centres. They found that fluid always moves through the gap separating the two cylinders but that eddies form for gap widths smaller than 0.0446
times the cylinder radius. This observation is also relevant to the study of suspension feeders. Interestingly, in the evolution of chain formation in planktonic diatoms,
colonies show a tendency to become more and more disjointed, while conserving appreciable stiffness (Beklemishev 1959). The most straightforward interpretation is
that cells thereby avoid each other’s depleted diffusional boundary layers and allow flow between cells.
Observations: nutrient uptake by motile cells in stagnant water

We know of remarkably few attempts to test these theories even in the engineering context. Empirical data obtained from electrochemical measurements for large
values of Pe are in good agreement (within 4%) with Equation 17 (Kutateladze et al. 1982). We are not aware of similar experiments for intermediate and low Péclet
numbers. Experiments with live organisms are even harder to conduct. While batch cultures are often maintained without agitation, the parameters needed to test the
theory are not measured. Furthermore, it is a fallacy to regard unstirred vessels as stagnant. It is difficult to avoid circulation due to thermal gradients without deliberate
water jacketing and prevention of evaporation, as anyone who has tried to measure slow settling velocities knows. Conversely, without stirring, cultures can become
inhomogeneous both in cell concentrations and in nutrient concentrations. It is thus difficult to make valid assumptions about velocity and concentration fields in typical
batch culture.
Nutrient fluxes on the scale of interest are difficult to detect. Therefore a common approach is to measure nutrient uptake by cells. Nutrient uptake is sensitive to
physiological state of the cell and depends on conditions under which cells have been grown (Dortch et al. 1991). It represents a lower bound on flux to the cell and
reflects an increase in flux quantitatively only so long as the uptake system is not saturated. Moreover, experimental errors may often be comparable in magnitude to the
predicted enhancement of flux, especially if test organisms are very small.
Testing the contribution of swimming and sinking to flux in laboratory settings is subject to added logistical difficulties. Canelli & Fuhs (1976) attempted to examine
the effect of
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sinking on phosphorous uptake by the diatom Thalassiosira fluviatilis by drawing an analogy with cells held on a filter through which a constant volumetric flow rate
of nutrient medium passed. The flow regime in this experiment cannot be described by Equations 16–18 and is more analogous to the flow of pore waters through
sediments (i.e. may be more relevant for diatoms experiencing pressuredriven flows in the benthos or in planktonic particle aggregates). This study demonstrated an
effect of advection, but its results cannot be related quantitatively to natural fluid motions on single cells in the water column.
Mass transfer to cells in steady shear flow
Theory
Nonmotile cells in steady shear flow

Another flow field that a cell can encounter is steady shear. Steady shear flow is associated with viscous dissipation of smallscale turbulence and with flows created in
shear tanks to study nutrient uptake by phytoplankton and bacteria. It also holds for viscous sublayers (e.g. bottom boundary layers). By definition, a steady shear flow
is one in which ∂U/∂t=0 and
(Equ. 23)
where V is the mean velocity vector, G=VU is the velocity gradient tensor (Gij=∂Ui/∂Xj) and X is a position vector (we use standard tensor notation, e.g. Spiegel
1959). From the point of view of a nonswimming, nonsinking planktonic osmotroph, only the relative velocity component (GX) is important. The velocity gradient (G)
can be decomposed into two components (Lighthill 1986), rotation rate (Ω=Ωij= 0.5(∂Ui/∂Xj–∂Uj/∂Xi)) and rate of strain (E=Eij=0.5(∂Ui/∂Xj+∂U/∂X.)):
(Equ. 24)
Unlike selfinduced rotation, which is typically parallel to the direction of motion, as in the case of dinoflagellates swimming in helical wave motion, a shear flow causes
the cell to rotate with an axis of rotation perpendicular to the velocity gradient (Fig. 4). While the straining motion of the flow acts to reduce the thickness of the
boundary layer around the cell and thus to enhance the flux of nutrients (Fig. 1), rotation will weaken the effect of advection on nutrient flux because it acts to close
streamlines, making them less effective in carrying nutrients to the cell (Batchelor 1979). Thus in a shear flow, the net contribution of advection will be determined by the
ratio |Ω|/|E|.
The governing equation for the concentration distribution around a cell in a linear, steady shear flow is again the dimensionless Equation 11 with the boundary
conditions (12, 13), where U* is the nondimensional velocity field of the shear flow. We define the characteristic velocity of the shear flow (|U|shear), and hence the
Péclet number of the cell, in terms of the shear rate E:
(Equ. 25)
(Equ. 26)
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Figure 4 Rotation of cells due to swimming and fluid shear. While cells swimming with a flagellum in a helical wave motion tend to
rotate so that their axis of rotation parallel the direction of translation, nonmotile cells suspended in a simple shear flow rotate
with their axis of rotation perpendicular to the flow direction. In the latter case rotation will weaken the effect of shearing on
nutrient flux relative to a nonrotating cell in shear. Passive, flowinduced rotation causes streamlines around the cell to curve
or even close, making them less effective in carrying nutrients.
where the shear rate is defined as
(Equ. 27)
The dimensions of a shear rate [T−1] can be easily intuited by noting that a shear is a difference in velocity [LT−1] across a distance [L]; division eliminates the length
scale and yields the rate. Its magnitude can be estimated from measurements of kinetic energy dissipation rate of the flow (ε, [L2 T−3]) since for any given shear flow
(Equ. 28)
and thus
(Equ. 29)
where v is the kinematic viscosity (L2 T−1). The characteristic length scale (Lc) is taken again to be the cell radius (r0).
The simplest shear flow is steady with all components of the rate of strain tensor (Eij)
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equal to zero except E12=E21=Ω=(γ/√2) (e.g. U=(γy, 0, 0); Leal 1992). This flow is not applicable directly to the upper mixed layer in the field but is of interest
because many laboratory experiments on the effects of shear on planktonic microorganisms have been performed in Couette devices that produce such shear flows
and because laminar sublayers of the boundary layers above the sea bed and surrounding objects in the sea do have this structure. A correlation between Sh and Pe for
onedimensional, steady shear flow was obtained by Frankel & Acrivos (1968) (see also Leal 1992) for Pe 1:
(Equ. 30)
After transforming to our previously defined E and Peshear,
(Equ. 31)
Because the axis of flowinduced rotation is perpendicular to flow direction, at high Péclet number—when rotation is strong and closed streamlines are formed around
the cell—Sh asymptotically approaches 4.5 as Pe ∞ (Frankel & Acrivos 1968).
In nature, however, flow experienced by very small cells is a threedimensional linear shear field whose orientation relative to the cell shifts continually. Solutions for
mass transfer in more general, steady, linear shear were derived by Batchelor (1979) for limiting cases. For Pe 1 and any pure straining motion (Ω=0):
(Equ. 32)
where the numerical coefficient is accurate within 3 %. The essence of pure straining motion at the scale of the cell is that the net hydrodynamic force acting on the cell
is zero, i.e. there is relative motion between the cell surface and the fluid but there is no net force that causes the cell to rotate or translate. When the ratio |Ω| /|E| ≤ 1
and Pe<<1,
(Equ. 33)
where the error in the numerical coefficient does not exceed 10%. When the ratio |Ω| /|E| > 1 and Pe<<1, only shear rate in the direction parallel to the axis of rotation
(Eω) contributes to advective flux:
(Equ. 34)
where we use Batchelor’s (1979) assumption that in the expression for E2. Thus it should provide a lower bound on Sh.
For Pe 1 and pure straining motion, the relation between Sh and Pe becomes (within an error of ≤1%)
(Equ. 35)
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Figure 5 Sherwood number (Sh) as a function of Péclet number (Pe) for nonmotile cells suspended in a steady, linear shear. Solid lines
are based on analytic solutions derived for Pe<<1 and Pe>>1. Equations 32 and 34 were derived for cells in linear shear in the
absence of rotation and in the presence of strong rotation, respectively, for Pe<<1 (Batchelor 1979). At small values of Pe
molecular diffusion is the governing transport mechanism, and rotation will have no significant inhibitory effect on advective
flux. At large Pe, however, strong rotation partially suppresses advective flux as indicated by the difference between the
results of Equation 35, which was derived for the nonrotating case and Equation 36, which was derived for strong rotation
(both by Batchelor 1979). Sh values for 0.1<Pe<90 were obtained numerically for the case of uniaxial, extensional flow (filled
circles; Appendix II). The curve fitted to the numerical values (for 0.1 Pe<90) is of the form Sh=0.63Pe0.4+0.82 (not shown).
Based on the scheme used for the model, we estimate the absolute error in our numerical solution for small Pe to be <0.005.
For large Pe the estimated error is predicted to be <1% of Sh. We are not aware of any general analytic or numerical solutions
for the region 0.01 <Pe<100, in which many marine osmotrophs fall. Therefore, estimates of Sh for that region were obtained
by interpolation for each of the two cases (i.e. pure straining motion and shearing motion in the presence of strong rotation).
Interpolations assume a function of the form Sh=a+bPec, where c for each case falls between the limits of its values for the
interpolation through the lowPe solution at Pe=0.01 (Sh=1.004+0.32Pe1/2 in the absence of rotation (lower dashed line);
Sh=1.002+0.21Pe1/2 in the presence of strong rotation (lower dotted line)) and the high Pe solution at Pe=100
(Sh=0.883+0.71Pe1/3 in the absence of rotation (upper dashed line); Sh= 0.921 +0.47Pe1/3 in the presence of rotation (upper
dotted line)). The region between these two interpolation limits is marked in gray. Open circles denote the empirical data of
Purcell (1978), who studied heat transport from spheres in a steady, twodimensional, pure straining motion (i.e. no rotation).
Explanations for the discrepancies between the analytical results, our numerical model for high Pe values and Purcell’s results
are given in the text.
For Pe>>1 when rotation and strain both act,
(Equ. 36)
again making the same assumption about the relation between Eω and E. Equations 35 and 36 demonstrate the relative reduction in mass transfer owing to rotation at
high Pe (Fig. 5). We
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are not aware of published analytic or numeric solutions for intermediate values of Pe; therefore for the region 0.01<Pe<100 our plotted values for Sh were obtained
by interpolation through the lowPe solution at Pe=0.01 and the highPe solution at Pe=100 (Fig. 5). Our interpolations assume conservative functional form, i.e.
Sh=a+bPec, where c falls between the limits of its values for high and low Pe and a and b are chosen to fit the two endpoints. We also obtained Sh values for the
region 0.1 <Pe< 90 by solving the problem numerically for the case of pure straining motion (Figure 5; Appendix II). When rotation is not zero, the problem becomes
threedimensional, and we do not attempt even numerical solution here. For 0.1 <Pe< 10 our numerical results fall in the region between the two interpolation limits
obtained for purely straining flow (Fig. 5). For larger Pe, Sh values predicted by our numerical model are higher than the upper interpolation limit. One explanation for
this discrepancy is that Equation 35 provides only a firstorder approximation for the increase in flux. We are not aware of higherorder expansions for the case of
shear flow but we expect that such solutions will include terms of order smaller than Pe1/3 (e.g. a constant, as in the case of the higherorder expansion for uniform flow
(Equation 17)). A function of the form Sh=0.9Pe1/3+c was fitted to our numerical results in the region 50<Pe< 90, suggesting that a constant of 0.56 should be added
to Equation 35.
Laboratory experiments
Validation of the theory in an engineering context

Measurements of mass transfer in shear flow are consistent with theory for large Pe (i.e. Sh Pe1/3; Kutateladze et al. 1982). The value of the proportionality constant
suggested by the data, however, is 10% smaller than suggested by Equation 35. This discrepancy is not surprising since the experimental setup (fixed sphere in a shear
flow) did not exactly mimic the case of a sphere suspended in an ambient shear flow for which Equation 35 was derived (Kutateladze et al. 1982).
Empirical results that include the region of intermediate Pe (Fig. 5) were provided by Purcell (1978), who performed an experiment to study the transport of heat
from a spherical particle suspended in a steady, twodimensional, purely straining shear (i.e. no rotation). The sphere in his experiments was immersed in a fluid of high
viscosity, and the velocity field was established by four counterrotating cylinders. From the temperature difference between the surface of the heated sphere and the
fluid, he obtained a relationship between heat transfer and shear. Purcell’s results indicate that, for Pe 1, Sh is proportional to Pe2. This disagreement between theory
and experimental results may be the result of the sensitivity of mass transfer at small Pe to flow geometry at large distances from the body. Shear in Purcell’s experiment
was disturbed by boundaries about eight radii from the sphere, while the analytic solution considers an unbounded flow field (Batchelor 1979). Nevertheless, Purcell’s
results have been used by several authors to predict the effect of turbulence on fluxes of nutrients to suspended osmotrophs (Lazier & Mann 1989; Jumars et al. 1993).
We (Jumars et al. 1993) compounded confusion by finding good fit of Purcell’s (1978) data to an inappropriate equation, i.e. one derived for Stokes’ flow past a
sphere (Clift et al. 1978; their equation 3–49), before we had encountered the full diversity of lowRe flow regimes of interest in questions of nutrient flux to and from
cells. We further failed to transform Clift et al.’s equation to our definitions of Pe and Sh.
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Nutrient uptake in steady shear

Very few experimental data are available for testing this theory in an oceanographic context. Logan & Dettmer (1989) investigated the effect of a laminar shear on
leucine uptake by bacteria for shear rates ranging from E=0s−1 to E≈56s−1 (0≤γ< 80s−1), although for shear rates of E>35s−1 the fluid became unstable and could no
longer be considered laminar. Shear rates were estimated from simple engineering relations for stirred vessels (Logan & Dettmer 1989, Van Duuren 1968). The model
for simple, steady, shear flow (Equation 31) predicts an increase of 3 % in leucine flux (Dleucine=7×10−6cm2s−1) for a cell with a radius of 0.45µm exposed to a shear
rate of 35s−1. Uptake by bacteria exposed to shear, however, did not increase compared with uptake by bacteria suspended in stagnant fluid. Instead, high shear rates
seemed to impair leucine uptake. Shear rates used in this experiment were far above any typical values encountered by bacteria in the ocean (shear rates in the mixed
layer varying from 0.01 to 1s−1), and the observed inhibition of uptake by shear may have resulted from stress or even mechanical damage.
In a later study, Logan & Kirchman (1991) investigated the effect of fluid shear (E<1.5s−1 for γ≤2.1s−1) on leucine uptake by natural assemblages of marine
bacteria. Their results showed no significant (less than 10% departure from unstirred vessels) enhancement of leucine uptake at shear rates 0<E<1. Their results agree
with the model prediction (Equation 31) that for organisms of typical bacterial size laminar shear is ineffective at increasing uptake of molecules with a diffusion
coefficient of the order 10−5–10−6cm2s−1. Their experimental error (standard deviation being 5% to 32% of the mean), however, is larger than the increase in the flux
predicted by Equation 31; therefore, the experiment cannot provide a strong test of the theory.
Few experimental data are available for larger organisms. Pasciak & Gavis (1975) studied =1.9×10−5cm2s−1, Li & Gregory 1974). The offset between
experimental results and theory may result from some combination of greater saturation of the uptake system under conditions of shear (predicted flux, therefore, over
estimating uptake), failure of the theory to include the shape effects of nonspherical cells and experimental error, whose magnitude was not provided. In summary,
there are no published data that show statistically significant departure from model predictions for simple shear flow, but no strong tests have been performed.
Mass transfer to cells in turbulent water
Nonmotile cells in turbulent flow

Turbulence acts to mix water and increase rates of momentum, heat and mass transfer. Turbulence is strongly intermittent in amplitude, space and time and is always
dissipative.
Page 92
Kinetic energy is transferred from large, turbulent eddies into smaller ones and then is dissipated as heat by viscous friction. The length scale of the smallest eddies
associated with the turbulent flow is known as the Kolmogorov length scale, η (Tennekes & Lumley 1972),
(Equ. 37)
where ε is the turbulent kinetic energy dissipation rate [L2 T−3] and v is the kinematic viscosity (approximately 0.01cm2s−2). Organized rotational motions of the fluid in
the absence of particles are thought not to exist below this scale. Typical values of the energy dissipation rate in the oceanic upper mixed layer range from 10−2 to
10−5cm2s−3 (Oakey & Elliott 1982, Gargett 1989, Brainerd & Gregg 1993). Thus, in the upper mixed layer the Kolmogorov scale is on the order of 1–6mm. Above
the Kolmogorov scale, the flow is turbulent (i.e. irregular and dominated by inertial forces), while below it viscosity dominates, resulting in laminar shear (Table 4).
Another feature of the turbulent flow field below the Kolmogorov length scale is that the statistical state of smallscale fluctuations is considered to be homogeneous,
isotropic and practically steady (Monin & Yaglom 1975).
Table 4 Characteristic flow fields at large distance from the cell, dominant transport mechanisms and proposed solutions for nutrient tranfer to cells.
Scalings Flow regimes Dominant transport mechanisms Relevant equations

Re « 1, r0 ŋb, Pe 1 Steady, laminar shear Molecular diffusion Equation 48
Re 1 , r0 ≈ ηb, Pe≈1 Steady, laminar shear Molecular diffusion and laminar advection No solution available (see Fig. 6)
Re 1, ηb<r0<η, Pe>1 Statistically steady, laminar shear Laminar advection No solution available (see Fig. 6)
Re<1, r0≤η, Pe 1 Statistically steady, laminar shear Laminar advection Equation 49
Re≈1, r0≈η, Pe 1 Transitional flow (between laminar and turbulent) Laminar and turbulent Outside the scope of this review. Do not
Re>1, r0>η, Pe 1 Turbulence Turbulent advection attempt to apply our equations
Hence, ambient flow in the vicinity of phytoplankton and bacteria (except for large chains of diatoms and filamentous cyanobacteria) can be very well approximated as
a linear shear field. Based on published field measurements of microstructure Lazier & Mann (1989) suggested that the smallest energycontaining eddies have a length
scale larger by a factor of 5–10 than the Kolmogorov scale and proposed that a coefficient of c. 10 should be used with the right side of Equation 37. A major
limitation in addressing the role of turbulence in the nutrition of planktonic osmotrophs is the rarity of flow measurements at microscopic levels. Current knowledge of
the flow regime in the vicinity of planktonic microorganisms is based solely on theoretical arguments and on measurements of flow at macroscopic levels. Accepting
Equation 37 as representing the size of the eddies below which the flow field can be well approximated as a laminar shear field will not result in a large error, however,
so long as the cells in question are much smaller than η produced by Equation 37.
A second relevant length scale addresses the smallest variations in the ambient concentration field. It was introduced by Batchelor (1952) but has seen far less
frequent application than Equation 37:
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(Equ. 38)
This length scale is smaller than the Kolmogorov scale because diffusion coefficients are smaller for molecules such as nutrients [O(10−5cm2s−1)] than for momentum
[O(10−2cm2s−1)]; mass transfer requires displacement of molecules, while momentum can be transferred in successive molecular collisions without as much net
displacement. This length scale separates the region in which the principal transport mechanism is shearing motion versus diffusion (Table 4). At length scales larger than
η, but smaller than η, temporal changes of the velocity field are slower than the Kolmogorov time scale of (v /ε)1/2, and the flow is considered to be statistically steady
(Table 4). Below this scale, molecular diffusion dominates transport and has a characteristic time scale (where Lc is a characteristic length scale). If we choose Lc
to be (vD2/ε)1/4, then the characteristic timescale of diffusion becomes (v/ε)1/2. This result implies that below ηb diffusional adjustment is faster than variations in the
flow, and the flow can therefore be approximated as steady shear. In the mixed layer, ηb may range between 32 μm and 180μm (assuming a kinetic energy dissipation
rate of 10−2 to 10−5cm2 s−3 and D=1×10−5).
A turbulent flow field is often decomposed into its mean (e.g. over an ensemble of measurements) and fluctuating components as
(Equ. 39)
The strain tensor can be decomposed similarly as
(Equ. 40)
Since in a turbulent flow
(Equ. 41)
(Tennekes & Lumley 1989), the relation between shear rate and kinetic energy dissipation rate is effectively
(Equ. 42)
For isotropic turbulence (eii=ejj, eij=ek l, i≠j, k≠l, and the mean rate of extension equals zero), only one component of the strain tensor is needed to describe the
relationship between ε and shear rate. The relationship between the r.m.s rate of extension in a fixed direction and ε was given by Taylor (1935),
(Equ. 43)
and,
(Equ. 44)
The local axis of rotation, however, changes over time. A relation between the mean rate of extension parallel to the local axis of rotation and ε was given by
Batchelor (1980),
(Equ. 45)
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where S is a skewness factor of the rate of extension in a fixed direction. Its values range from 0.3 to 1, based on measurements in various turbulent flows and over a
wide range of Reynolds numbers (Batchelor 1980). Since the measures for oceanic turbulence are ε and v, we define the characteristic velocity and hence Pe in terms
of those parameters:
(Equ. 46)
and
(Equ. 47)
We are not aware of solutions for mass transfer to particles suspended in turbulent flow for small or intermediate values of Pe. For cells smaller than ηb (which by
definition means Pe<1), however, the flow can be assumed to be steady. In addition, since in isotropic turbulence the ratio between Eturbulence and Ωturbulence is of
order 1, Equation 33 (steady shear flow at Pe 1, where E/Ω≤1) can be used to describe mass transfer to cells suspended in turbulent water. After replacing Peshear
with Peturbulence, using Equation 42 to convert E to terms of ε/v, Equation 33 becomes
(Equ. 48)
For cells larger than ηb but smaller than η, Equation 36 (for Pe 1) can be used after converting Eω to terms of ε and v, according to Equation 45 (Batchelor 1980).
Batchelor (1980) used S=0.6 to solve for mass transfer to spherical particles in statistically steady (but not necessarily homogeneous), turbulent flow:
(Equ. 49)
The leading coefficient in this equation will vary by 20% depending on the choice of S. We caution that varying definitions of shear rate (and hence Pe) have been used.
In several
Figure 6 Sherwood number (Sh) as a function of Péclet number (Pe) for nonmotile cells suspended in turbulent water. For Pe 1 (i.e.
cells smaller than ηb) we used Equation 48 for steady shear flow, and for Pe 1 we used Equation 49, which was derived for
statistically steady shear flow. For 0.01<Pe< 100, Sh values were obtained by interpolation (gray area) as described in Figure 5
where the lower limit for interpolation is given by Sh=1.014+0.15Pe1/2 and the upper limit is given by the line
Sh=0.955+0.344Pe1/3.
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studies, a less rigorously defined shear rate was related to 8 by an arbitrary leading coefficient of 0.5 (e.g. Bowen et al. 1993, Jumars et al. 1993). In the absence of
analytic or numeric solutions for intermediate values of Pe, Sh values used here for the region 0.01<Pe<100 were obtained by interpolation (Fig. 6).
Application to the ocean

Turbulent kinetic energy dissipation rate (ε) in the ocean shows great variability in both space and time, making the choice of a representative value of the shear
experienced by cells challenging and maybe unwise. Within the mixed layer, successive estimates of ε differ by an order of magnitude (Shay & Gregg 1986). Turbulent
kinetic energy dissipation rates are high at the surface and decay with depth, often decreasing by an order of magnitude per meter (Brainerd & Gregg 1993).
Dissipation rates vary daily, seasonally and annually. Furthermore, in the nearsurface zone, values of 8 obtained a few hours apart may differ by three orders of
magnitude (Brainerd & Gregg 1993). Turbulence can be beneficial if cells are suspended in high shear long enough to allow appreciable enhancement of nutrient flux.
Since published values of estimated turbulent kinetic energy dissipation rates are time averaged, one must know the distribution function of 8 in order to estimate the
fraction of the time that a cell is exposed to a given shear. The distribution function of dissipation estimates drawn from surface mixed layers is nearly lognormal (Osborn
& Lueck 1985, but see Yamazaki & Lueck 1992) and is parameterized by and s2Inε, the mean and the variance of lnε. The latter is a measure of the intermittency
of the turbulence. For example, taking a value of the variance in the upper mixed layer s2Inε=1.5 (Baker & Gibson 1987), the dissipation rate will have a value equal to
or smaller than the mean 73 % of the time (Shimeta 1993). Because of the strong intermittency of turbulence, its effect on flux will vary greatly in space and time and
will probably be more significant in coastal zones, where turbulence is more intense, than in the open ocean.
If we assume a characteristic value of ε in the surface layer to be 10−2cm2s−3, theory suggests that turbulence in the ocean will favour largecelled competitors. The
critical cell size for 50% increase in the flux (D=1×10−5cm2s−1) at this dissipation rate ranges between 63–100μm (Table 2). Planktonic bacteria (other than
filamentous bluegreens) and small flagellates are well below this limit, while large cells, chains and colonies are above it. These calculations indicate a stronger effect of
turbulence than previously supposed. Assuming a diffusion coefficient D=2×10−5cm2 s−1, Lazier & Mann (1989) calculated, based on the experimental results of
Purcell (1978), an increase in the flux of only 2% for cells 50μm in radius in the presence of strong turbulence (ε=10−2cm2sec−3)—while the model we use suggests an
increase a full order of magnitude greater (18–32%; Fig. 6).
The behaviour of cells in turbulent flow is not well known but is likely to vary strongly with shape. Since large cells, chains and colonies are the ones expected to
benefit most from turbulence, a better understanding of their behaviour in turbulent flow is needed. Whether chains are straight, bent, rigid or flexible will affect their
behaviour in shear flow and hence their nutrient fluxes and competitive abilities. For instance, the period of rotation of permanently bent, threadlike particles is
appreciably lower than that of straight particles (Forgacs & Mason 1959). If this disparity holds in the pelagic environment, one may expect straight chains to
experience reduced flux compared to bent chains. In both theoretical and empirical results Van de Ven & Mason (1976) showed that the behaviour of chains of equal
sized
Page 96
spheres in simple shear flow depends on whether the units in the chain touch each other. A straight chain of touching units will rotate as a rigid body when exposed to
shear while a chain of nontouching spheres will change length periodically with their orientation. Curved chains that are not rigid will bend. Although the units in chains
of nontouching spheres were not attached directly to each other by means of thin, flexible threads (as in the case of some diatom species), but were held together by
means of electrostatic and van der Waals forces, and although the results of their study are limited to Poiseuille flow and short chains (up to 5 units), they are intriguing
in the light of the diverse morphologies of chains observed in nature and the evolutionary trend in diatom chain formation toward nontouching cells. Details of spacing
within chains will alter the way that turbulence and bending “pump” nutrientreplete water between the cells of chains.
There are no measurements of nutrient uptake by cells suspended in controlled turbulence that can be tested against this theory. Results from experiments done in
bottles placed on shaker tables and plankton wheels or stirred with stir bars do not provide strong insight about turbulence effects since the nature of the flows and the
magnitudes of shear are neither well quantified nor related to nature. Savidge (1981), who investigated the effect of turbulence on uptake rate of phosphate and nitrate
by the diatom Phaeodactylum tricornutum, used an oscillating grid to produce turbulence. Unfortunately, neither shear rate nor energy dissipation rate was
determined, and no information was provided about sizes of cells or the magnitude of experimental error. Oscillatinggrid turbulence shows extreme spatial variability,
the dissipation rate falling off as the fourth power of distance from the grid (Brumley & Jirka 1987). Therefore, no quantitative conclusions can be drawn from this
experiment. Better indication at high Pe comes from chemical engineering experiments on dissolution rates of particles in turbulent, stirred flows. These data have been
reviewed and summarized by Batchelor (1980), who found good fit to the predictions.
Translational motion in turbulent water
Swimming and sinking in turbulent water

The solutions presented so far for mass transfer in the presence of turbulenceinduced shear assumed that the cells were suspended without translation in turbulent
water. In the real ocean, however, cells in turbulent water may swim or sink. The resultant flow field can be expressed as a superposition of steady translational motion
and shear (Batchelor 1980). Total nutrient transfer to motile cells, for large Pe, becomes (Batchelor 1980):
(Equ. 50)
The coefficient a depends on the ratio 5r0e11/Us* (where the asterisk indicates that only the component of the swimming or sinking velocity parallel to the axis of
rotation is used. It varies between 0.495 (when 5r0e11/Us* ∞, i.e. pure shearing motion of Equation 49) and 0.545 (when 5r0e11/Us*=0, i.e. pure translational
motion as given by the firstorder approximation of Equation 17). As mentioned previously, if a cell is rotating and closed streamlines are formed, translational motion
will contribute to the flux in proportion to the magnitude of its component in the direction parallel to the axis of rotation. If isotropic turbu
Page 97
lence is assumed, however, the cell is continually reoriented, and the mean value of the component of translational motion parallel to the axis of rotation equals zero.
Thus in turbulent water, as soon as the shearinduced rotation rate reaches a critical magnitude (Ω=UsPe−1/3/r0, where Pe=r0 Us/D), swimming or sinking will have no
appreciable effect on transfer of nutrients (Batchelor 1980). Even if the swimming mechanism imparts rotation parallel to the instantaneous swimming direction, high
turbulence intensities may overwhelm the rotational component and effectively randomize the direction of translation.
A common observation, both spatially and temporally, is that small, motile phytoplankton cells tend to predominate in stratified, calm, oligotrophic water while large
cells (mainly diatoms) prevail in turbulent, nutrientrich water (Margalef 1978, Malone 1980, Mann 1992). Dominance of flagellates in calm water is probably a result
of their ability to increase nutrient uptake through swimming—both from reaching layers of higher nutrient concentration and within any particular nutrient concentration
from relative motion. In turbulent water, on the other hand, dinoflagellates may not be able to maintain a rotational axis parallel to the direction of the swimming or to the
direction of the shear flow; thus they will suffer a relative reduction in nutrient flux. Predicted critical velocities below which swimming is no longer effective in enhancing
the flux are of the same order of magnitude as maximal swimming velocities measured for dinoflagellates (Figure 7). Thus swimming, especially for large dinoflagellate
cells, may not contribute significantly to flux under natural turbulence. There is increasing evidence that dinoflagellates are sensitive to strong shear. Under conditions of
vigorous turbulence cells may suffer mechanical damage. Loss of flagella, changes in swimming behaviour, and growthrate inhibition have been observed for several
species of dinoflagellates exposed to straining motion in the laboratory (Thomas & Gibson 1990 a, b, 1992, Berdalet 1992).
Figure 7 Critical velocities below which swimming or sinking cannot significantly affect the flux in turbulent water, as a function of cell
size. Critical velocities were calculated based on Batchelor (1980) Ucritical=(r2/√D)(ε/v)3/4, with energy dissipation rate (ε)
taken to be 1×10−2cm2s−3, D to be 1×10−5 cm2s−1 and v to be 1×10−2cm2s−1. For cells swimming or sinking more slowly
(grey area under the solid line) turbulence so dominates the advective contribution to the flux that swimming or sinking is
ineffectual.
Page 98
Flowinduced translational motion

Cells and chains of sizes near the Kolmogorov scale may exhibit passive translational motion due to a lift force induced by curvature of the flow. Passive migration can
increase total mass transfer because it adds an additional component of relative velocity between the cell and its surrounding water and facilitates encounter of cells with
“new” water (i.e. changes C∞ ). The idea of translational motion induced by turbulence stems from observations of the migration of rigid, neutrally buoyant particles in
Poiseuille flow (a nonlinear shear flow with ∂ui/∂x j ≠ constant, i≠j, such as in pipes and blood vessels) at particle Reynolds numbers as low as 10−3 (Serge &
Silbergberg 1961, Karnis et al. 1963). Nonlinearity of the flow due to wall effects and small but nonnegligible inertial forces (Goldsmith & Mason 1967, Leal 1980)
impart the lift force that causes migration. Flexible particles show similar behaviour even at smaller Reynolds numbers as a result of lift due to shear deformation
(Goldsmith & Mason 1961, Karnis et al. 1963). The direction of the migration of neutrally buoyant particles in Poiseuille flow is to the region of minimal shear, while the
direction of migration of nonneutrally buoyant particles depends on whether the particles lead or lag the local undisturbed motion (Leal 1980). Since a similar
phenomenon has also been predicted in unbounded flows (Saffman 1965a) it may be relevant for large cells in turbulent water.
To calculate the translational velocity that results from curvature of the flow, we assume a onedimensional flow field of one Fourier component with the length scale
of the smallest energetic eddy. This simplification is suitable for our purpose since the turbulent flow field is assumed to be isotropic. We assume a freestream flow
velocity,
(Equ. 51)
where is the Kolmogorov scale of velocity ((εv)1/4), η is the Kolmogorov length scale, and y is the spatial variable. Transverse (to the streamline) velocity of the
phytoplankter due to lift is (Saffman 1965a, b, McLaughlin 1991)
(Equ. 52)
where Reshear is the Reynolds number of the particle based on the turbulenceinduced shear and is defined as
(Equ. 53)
Equation 52 is independent of rotation, and the error is of order Re 1. ΔU is the relative velocity between the flow speed at the centre of the particle (when the
particle is removed) and the speed of the particle. The difference in the flow velocity and the particle velocity arises from the nonlinearity of the shear flow. Equation 52
holds only when the cell is not motile or if Re based on its swimming or sinking velocity (Res) is much smaller than the square root of the cell Reynolds number based on
shear velocity (Reshear). McLaughlin (1991) expanded Saffman’s analysis to cases in which Res≥√Reshear and demonstrated that migration velocity increases as the
ratio √Reshear/Res increases. If we assume a typical shear rate for strong turbulence to be c. 0.5 s−1, for a cell 50μm in radius Reshear becomes O(10−2). Re for
sinking cells is typically smaller than 10−2 (Table 1), and Equation 52 can be used to
Page 99
calculate the translational velocity due to flow curvature. Re for swimming cells can be larger, and the corrections to Equation 49 suggested by McLaughlin (1991, his
table 1) should be used.
In our case, the r.m.s. relative velocity that arises from the curvature of the flow is the second term of the Taylor expansion for particle velocity:
(Equ. 54)
Thus,
(Equ. 55)
or
(Equ 56)
For an organism 50–100μm in radius in turbulent water with ε=O(10−2)cm2s−3 the transverse velocity is about 10−2–10−4 body lengths s−1. The direct contribution of
shearinduced lift to the advective flux will strongly depend on cell size and will become important only when cell size is of order η. Our calculations suggest that
migration due to curvature of the flow is potentially important in laboratory experiments but is probably of limited relevance to the pelagic environment.
Conclusions and prospects

Nutrient fluxes to planktonic osmotrophs in the presence of fluid motion depend on three sets of physical characteristics: the flow regime, the size and shape of the cell
and the diffusion coefficient of the nutrient of interest. Solutions for the advective contribution to flux clearly do not carry over from one flow regime to another. Careful
attention should be given, therefore, when comparing theoretical results with experimental or field data. For example, theoretical predictions derived for a cell in three
dimensional, isotropic turbulence cannot be tested directly by experiments in Couette flow devices. Advective flux in a given flow field will also be affected by the
behaviour of the cell, e.g. rotation or deformation of flexible chains. Direct observations of behaviours of cells in shear flows or even of behaviours of cells while
sedimenting, however, are practically nonexistent.
A recurring theme is the potential effect of rotation on flux for all but one orientation, i.e. rotation about an axis parallel to the flow. We are not aware of any
measurements of the magnitude of the rotation effect at any Pe. It could be explored in stagnant water and linear shear with slight modification of the approach of
Purcell (1978) and seems important to an understanding of the ecology of dinoflagellates in particular. A key question is whether dinoflagellates in the presence of
steady shear of varying magnitudes or of even gentle turbulence can maintain a rotational orientation that does not hinder nutrient exchange. It seems likely, however,
that dinoflagellates are ill suited for experimentation in the Couette devices that are typically used to mimic some turbulence effects. At low shears their tendencies to
Page 100
swim in more or less straight lines will cause them to interact frequently with the walls, and at high Pe they should suffer reduced fluxes from rotation for any rotational
orientation that is not perfectly aligned with the curved plane of shear.
The contribution of advection to nutrient flux will increase with cell size. For nutrients of small molecular sizes, such as nitrate, phosphate, glucose or individual amino
acids, fluid motion will have small effect on the flux to small planktonic organisms, such as bacteria, but will be important for large phytoplankton cells and osmotrophic
larvae. Chains and filaments, in particular, are expected to experience enhanced relative motion if their lengths approach the Kolmogorov scale. Fluid motion can be
very significant, however, to small cells that feed on large dissolved molecules or colloids as implied from the inverse dependence of Sh on the diffusion coefficient. The
dependence of Sh on cell size differs among the mechanisms by which relative motion can be induced (Table 3). Hence, optimal cell size will vary with this changing
dependence (Jumars et al. 1993). This result may provide one explanation for the existence of such a wide range of phytoplankton sizes. While the theory that relates
cell size and mass transfer is well developed, there is clear need for more laboratory experiments. The size range that has been examined so far is very narrow and
limited to small cells, mainly bacteria.
A striking feature of the phytoplankton community is its diversity of shapes and morphologies. Analytic solutions for mass transfer to shapes other than spheres are
limited to spheroids and cylinders. Solutions for more complex shapes require numerical modelling. It has been suggested that spines and horns, common features in
many phytoplankton species, are devices that increase drag and thereby decrease sinking speeds or are means to thwart grazers (Sournia 1982). An additional function
of horns and spines may be to increase the effective size of cells without changing volume or gross catabolic rate, so that absolute length will exceed the Kolmogorov
scale or at least Batchelor’s (1952) diffusional length scale (Table 4). Chain formation can serve a similar purpose, especially if cells are held apart from each other so
that their diffusionally depleted layers do not overlap. While cells in a chain can remain as small units, the total length of the chain may be long enough to experience the
curvature of a turbulent flow and gain benefit from the relative difference in velocity. Terminal cells would experience the greatest relative motion, and chain flexibility
would influence absolute amounts of motion experienced and their time variation.
Although most osmotrophs live at low Reynolds numbers with viscous forces smoothing out velocity gradients, the potential flow fields experienced by cells are
varied, and comparably diverse strategies can be used by cells of various species to alter their physical environments and hence their diffusional fluxes. In such an
unsteady environment as the upper mixed layer there is no one strategy that will maximize nutrient transfer to the cell at all times or for all cell sizes.
Acknowledgments
We thank K.Banse, H.Berg, T.Daniel, N.Franzen, G.Jackson, J.Lazier, B.Logan, K. Mann, A.R.M.Nowell, M.J.Perry, J.Shimeta and P.Yager for constructive
comments on early versions of the manuscript. We are grateful to H.Berg for reexamining and sharing the details of his numerical model. LuAnne Thompson kindly
provided her “big machine” for running our numerical model. This study was supported by ONR Grants N00014–94–1– 0656 and N00014–94–0264.
Page 101
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Appendix I Notation
Symbol Definition Dimensions

A cell surface L2
C concentration of a given nutrient mol L−3
C∞ ambient concentration, far from the cell surface mol L−3
C0 concentration at the cell surface mol L−3
C* dimensionless concentration (C/C∞ ) dimensionless
D diffusion coefficient L2T−1

E rate of strain tensor T−1
E shear rate E |E| T−1
E the ij component of the strain rate tensor T−1
ij
Ē mean shear rate T−1

Eω component of the strain rate tensor parallel to the direction of rotation T−1
e perturbation strain rate T−1
eij the ij component of the perturbation strain rate tensor T−1
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f nondimentional drag on the cell dimensionless
G velocity gradient tensor T−1
G the ij component of the velocity gradient tensor T−1
ij
g gravitational acceleration LT−2
k mass transfer coefficient LT−1
K
m
concentration at which uptake equals 1/2Vmax molL−3
Lc characteristic length scale L
n unit vector inward normal to the body surface dimensionless

P ratio between diffusion and maximal uptake rates (Pasciak & Gavis 1974) dimensionless
Pe dimensionless
Péclet number,
Peshear dimensionless
Péclet number in shear flow,
Peturbulence dimensionless
Péclet number in turbulence,
Q total flux arriving to the cell in the presence of fluid motion mol cell−1 T−1
QD purely diffusional flux mol cell−1 T−1
r radial distance from the centre of the cell L
r0 cell radius L
r * nondimensional radial distance from the centre of the cell dimensionless

Re dimensionless
Reynolds number. The ratio between inertial and viscous forces,
Res dimensionless
Reynolds number of a cell based on its swimming velocity,
Reshear dimensionless
Reynolds number of a cell based on the shear flow,
Sh dimensionless
Sherwood number,
Sh0 Sherwood number in the case of pure diffusion dimensionless
tD characteristic time scale for diffusion T
U characteristic velocity (U=U1, U2, U3) LT−1

U velocity field LT−1
U* dimensionless velocity field (U/U) dimensionless
Ū mean velocity LT−1
U∞ farfield velocity LT−1
Usinking sinking velocity LT−1
Uswimming swimming velocity LT−1
Us swimming or sinking velocity LT−1
Us* Component of swimming or sinking velocity parallel to the axis of rotation LT−1
Ushear characteristic velocity of the shear flow LT−1
Utransverse velocity of passive migration LT−1
Uturbulence characteristic velocity of subKolmogorov turbulent flow LT−1
u′ perturbation velocity LT−1
V mean free velocity of the shear flow LT−1
Vmax maximal uptake rate mol cell−1 T−1
X position vector X=(x1, x2, x3)=(x, y, z) −
ε kinetic energy dissipation rate of the flow L2T−3

ŋ Kolmogorov length scale (of the smallest eddies associated with the shear flow) L
ηb Batchelor length scale (of the smallest fluctuations in the concentration field) L
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γ shear rate for simple shear flow T−1
del operator that indicates partial derivatives in three spatial dimensions; L−1
in Cartesian coordinates,
* dimensionless del operator dimensionless
µ dynamic viscosity ML−1 T−1
υ kinematic viscosity L2T−1
Ω rotation rate tensor T−1
Ωij T−1
the ij component of the rotation rate tensor,
ρ density of the medium in which cells are immersed ML−3
ρc mean density of the cell ML−3
υ Kolmogorov scale of velocity LT−1
Appendix 2 Advectiondiffusion model

We solved numerically for concentration distributions of nutrients around a spherical cell at low Reynolds number under unidirectional flow and linear shear (Re 1).
The concentration field around the cell for both flow regimes is given by Equation 7 with the boundary conditions 8 and 9, where, for spherical coordinates,
and
The velocity field U depends on the flow regime. For unidirectional, uniform flow past a sphere, U is composed of two components, Ur and Uθ, where (Leal 1992)
Since the problem is axisymmetric, velocity and concentration do not depend on the second angular coordinate, The velocity field of the shear flow
(uniaxial extensional flow where E=(2,0,0; 0, −1, 0; 0,0,−1) also comprises two components, Ur and Uθ, where (Leal 1992)
For each flow field, we nondimensionalized by cell radius, free velocity (far from the cell) and concentration at the domain boundary. Due to symmetry, only onehalf
the domain was
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computed, with an additional noflux boundary condition at the angular boundary (∂C/∂θ=0 at θ=0, π). In order to increase resolution near the cell the radial
independent variable r was replaced by the variable μ=ln(r). The grid (μθ coordinates) was composed of 231 points with constant radial separation (Δμ) and 101
points with constant angular separation (Δθ). Computation of the concentration at each point was marched in time using upwind difference. The firstorder spatial
derivative was calculated from central differencing and the secondorder spatial derivative was calculated using ADI (alternating direction implicit) following Press et al.
(1992, Ch. 19). The model was run until satisfactory steady state was reached, i.e. when the change in Sh was smaller than 10−4 for one nondimensional time step
(r02/D). Boundary conditions 9 cannot be applied as given in the text since r=∞ is not part of the domain. Therefore we used C(rmax)=1−(1/rmax), the value obtained
from the analytic solution for pure diffusion. The solution for the problem with this boundary condition will converge to the exact solution as grid size increases toward
infinity. For small Pe we chose the domain boundary to be O(100r0) and tested model sensitivity to it by doubling domain size. For large Pe values the domain we
needed was of O(1000r0) since the effect of the cell on the concentration field extends to a larger distance. Berg & Purcell (1977) used a similar model but with a
smaller domain (H. Berg, pers. comm.). Our sensitivity analysis with our own model suggests that their domain might not extend far enough to include the flow variation
contained in Stokes’ solution. We believe that this truncation of the flow and concentration field is the principal reason for the discrepancy between their model and the
asymptotic analytic solutions and our numerical model.
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FACTORS AFFECTING GRAZEREPIPHYTE INTERACTIONS IN

TEMPERATE SEAGRASS MEADOWS
P.JERNAKOFF, A.BREARLEY,1 J.NIELSEN2
CSIRO Division of Fisheries, PO Box 20, North Beach, WA 6020, Australia
1 Current Address: Botany Department, University of Western Australia, Nedlands, WA 6009, Australia
2 Current Address: Western Australia Museum, Perth, WA 6000, Australia

Abstract This review examines the interactions of grazers and epiphytes in temperate seagrass meadows. These interactions are critical to the dynamic
response of seagrass communities that are exposed to a range of internal and external pressures including anthropogenic eutrophication. To predict
successfully the outcome of grazerepiphyte interactions and the resulting impact on the seagrass community, a detailed knowledge of the key
processes is required, as well as a knowledge of the magnitude and direction in which they operate. Despite extensive study, the interactions of
seagrasses, grazers and epiphytes are poorly known and there have been several reviews on aspects of the research. We extend the scope of previous
reviews in a number of areas, particularly with respect to how differences in the lifehistory patterns of species interact at a variety of spatial and
temporal scales. We also review the implications of spatial and temporal variability of grazerepiphyte interactions to interpretations for studies carried
out at a single time or location.
The review is in three parts. The first reviews the biology of the main components of seagrass meadows that are relevant to grazerepiphyte
interactions (i.e. seagrasses, epiphytes and grazers). It examines the range of differences within each of the components that affect the outcome of
these interactions (e.g. differences between seagrass species in the size, shape and turnover time of leaves). The second reviews the spatial and
temporal scales at which interactions occur. Spatial scales range from within and between seagrass leaves to environmental gradients whereas
temporal scales vary from diurnal to annual. The third part reviews how interactions between grazers and epiphytes result in the overall pattern of
distribution and abundance. It reviews the implications arising from the scale at which interactions are measured in observational and experimental
studies. To predict the impact of grazerepiphyte interactions, one needs a detailed knowledge of the main processes taking place on several spatial
and temporal scales. Furthermore, one cannot simply extrapolate the results of studies from one location or time. The influence of localized conditions,
such as the species and morphology of the seagrass, the life history and recruitment patterns of epiphytes and grazers, and the physical environment
and its seasonal changes must all be taken into account.
Further research on grazerepiphyte interactions is needed before their role in seagrass communities can be accurately and reliably assessed. We
need a better understanding of the recruitment dynamics and dietary preferences of grazers (including assessment of the quality of food), of the local
factors likely to affect interactions, and of the likely magnitude of heterogeneity at a range of spatial and temporal scales.
Introduction
The interactions of seagrasses, grazers, and seagrass epiphytes are complex and still poorly understood despite extensive study (Robertson & Mann 1982, Orth & van
Montfrans 1984, van Montfrans et al. 1984, Howard et al. 1989, Brawley 1992, Mazzella et al. 1992, Orth 1992, Neckles et al. 1993). This complexity arises
because many factors act directly or indi
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rectly (or both) on the plant and animal components of the seagrassgrazerepiphyte community (Orth 1992, Neckles et al. 1993, Williams & Ruckelshaus 1993).
There have been several reviews of research on grazerepiphyte interactions in seagrass meadows (e.g. Orth & van Montfrans 1984, Mazzella et al. 1992, Orth
1992). Orth & van Montfrans (1984) give a comprehensive account of some of the factors affecting the interactions of plants and animals but much material has been
published since their review. Later reviews concentrated on particular aspects of these interactions. For example, Mazzella et al. (1992) concentrate on trophic
relationships in Posidonia oceanica ecosystems, while Orth (1992) gives a general overview of the physical and biological determinants of plantanimal interactions in
seagrass meadows.
We extend the scope of previous reviews in a number of areas. We examine the variability of seagrasses, epiphytes and their grazers at several spatial and temporal
scales and assess how these patterns affect grazerepiphyte interactions, with consequent implications for the interpretation of studies that are carried out at a single
location at one time of the year. We also explore how differences in the implications of different life history patterns (e.g. differences in the feeding mode of grazers) and
physical morphologies (e.g. differences in shoot density of seagrasses) affect the interactions.
Our approach is first, to review what is known about the general biology of the main components (i.e. seagrasses, epiphytes and grazers). Secondly, we examine the
patterns of epiphyte and grazer abundance and thirdly, we deal with the interactions themselves. Finally, we synthesize our findings to identify gaps in our knowledge of
the structure and linkages in key ecological processes within the seagrassepiphytegrazer community.
We have focused this review on temperate seagrass communities, with particular reference to southwestern Australia, which has a greater diversity of seagrass
species than any other region. However, we also discuss some research on tropical seagrasses to highlight where they resemble and where they differ from temperate
communities.
Much of the information is summarized in tables: ecological and morphological characteristics of seagrasses; studies of grazers of seagrass epiphytes; and a
comparison of the grazing rates of different taxa. The tables synthesize the results and conclusions of these studies and review the scientific methods that are used to
investigate the main components of seagrass meadows, and grazerepiphyte interactions themselves.
General biology
Seagrasses
Seagrasses are aquatic angiosperms that are generally restricted to soft sediment habitats. Seagrass leaves and stems provide the primary substratum for the attachment
of epiphytes within seagrass meadows and they are the basic structural component in which grazerepiphyte interactions occur. Seagrasses occur in a range of
environmental conditions, from the tropics through to cool temperate seas. Currently 58 seagrass species belonging to 12 genera, four families, and two orders are
recognized throughout the world. Of these, 30 species in 11 genera occur in Australia; Phyllospadix species are restricted to the northern Pacific (Kuo & McComb
1989).
On a regional scale, Western Australia has the highest diversity of seagrass species in the world (10 genera with 25 species) and some of the most extensive seagrass
beds (Kirkman &
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Walker 1989). The seagrasses there grow in both enclosed, relatively calm estuaries, with variable salinities and temperatures, and in exposed and turbulent nearshore
coastal areas (Kirkman 1985, Kirkman & Walker 1989, Larkum et al. 1989, West et al. 1989). The high diversity in this region has yet to be adequately explained but
has been attributed to the high level of endemism, the range of temperate and tropical habitats available for colonization, and the temperature and direction of prevailing
water currents (Kirkman & Walker 1989, Larkum & den Hartog 1989).
On a local scale, the patchwork of species within a multispecies meadow is considered to be a response to variable exposure to local conditions (Cambridge 1975,
Kirkman 1985, Fonseca & Kenworthy 1987). There are many different shapes and sizes of seagrasses. Their growth, distribution and patch dynamics are influenced
by environmental factors such as light (Dennison 1987), temperature (Bulthuis 1987), salinity (Walker et al. 1988, Hillman et al. 1989), water movement (Fonseca &
Kenworthy 1987) and nutrients (Bulthuis & Woelkerling 1981, Short 1987). The influence of these factors varies with the size and age of the seagrass patches (Olesen
& SandJensen 1994). The morphological characteristics and attributes of a species, including surface area, lifespan, seasonality of growth, and leaf morphology
(Table 1), affect the space available for epiphyte colonization, the degree of protection for algal propagules and the lifespan of the epiphytes.
A morphological classification developed by Borowitzka & Lethbridge (1989), based on den Hartog (1977), is useful when considering colonization of seagrasses
by epiphytes. These authors divide seagrasses into five groups:
(1) Species with long, subulate leaves, e.g. Syringodium;
(2) Species with fine, linear leaves, e.g. Halodule, Zostera subgenus Zosterella;
(3) Species with wide linear or strapshaped leaves, e.g. Cymodocea, Zostera subgenus Zosterella, Thalassia, Enhalus, Posidonia, Phyllospadix. (Referred to as
zosteroid types in this review);
(4) Species with delicate elliptic, ovate, lanceolate or linear leaves, e.g. Halophila;
(5) Species with upright, lignified stems and strapshaped leaves arranged distichously along them, e.g. Amphibolis, Thalassodendron, Heterozostera. (Referred to
as amphiboloid types in this review). Although Borowitzka & Lethbridge (1989) place Heterozostera in this group, it loses its reproductive stems and has a shorter
lifespan than Amphibolis and Thalassodendron. We suggest Heterozostera might be better placed under group (2).
Some of these morphological groups can be further divided according to the subhabitats they provide for epiphytes. For example, Amphibolis provides three main
subhabitats: the longlived, erect stem; the shorterlived, strapshaped leaves; and the cryptic habitat at the bases of leaf clusters (Borowitzka & Lethbridge 1989).
Borowitzka et al. (1990) record a greater abundance of epiphytic algae on the longerlived stems of Amphibolis than on the leaves.
Leaf size may affect the type and abundance of epiphytes. The size of a macroalga’s basal attachment disc might limit its ability to colonize macrophytes (den Hartog
1972). However, Kendrick et al. (1988) found Sargassum spp., which has a large basal disk, reproducing as an epiphyte on the seagrass Amphibolis antarctica.
Because algae with smaller discs can theoretically colonize a greater variety of macrophytes than those with larger discs, widerleaved macrophytes should support a
greater diversity of algae than narrow leaved forms. May et al. (1978) found a greater diversity of epiphytes on Posidonia australis with a large surface area than on
Zostera capricorni and Heterozostera tasmanica with less surface area. However, differences in the diversity of epiphytes might be a function of seagrass species,
loca
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Table 1 Ecological and morphological characteristics of seagrasses referred to in the review. *
Genus Species Distribution Habitat Temperature Depth Salinity (%o) Leaf Leaf Leaf No. of
(°C) longevity length width leaves
(days)* (cm) (mm)
Cymodoceaceae filiforme (as Caribbean, Gulf coastal tropical sublittoral MLWS polyhaline, 18 10–30 0.8–2 2–3
Syringodium Cymodocea of Mexico to 25 m euryhaline (12– circular
filiforme in text) 14)
Halodule beaudettei Caribbean, Gulf coastal sand and tropical low intertidal euryhaline, – 5–20 0.15–1.5 2
of Mexico mud polyhaline and
euhaline
Halodule uninervis N. Australia, bays with sand and tropical littoral, sublittoral euryhaline to – 6–15 0.5–3.5 2
Indian Ocean mud to 30 m hyperhaline
and West
Pacific
Halodule wrightii Caribbean, Gulf bays with sand and tropical lower eulittoral euryhaline 34 5–18 0.3–1 2
of Mexico mud and upper
sublittoral
Cymodocea serrulata Indian Ocean marine, sandy mud tropical mainly below low sensitive to a – 6–15 4–9 2–7
and West and coral sand water at spring decrease in
Pacific tides salinity
Amphibolis antarctica Southern near coastal and temperate sublittoral to 23 m (36–62.4) 28–40 18–30 3.5–10 8–10
Australia marine embayments
Amphibolis griffithii Southern near coastal and temperate sublittoral to 40 m stenohaline – 3–10 3–7 3–5
Australia marine embayments
Thalassodendron pachyrhizum Southern marine exposed rock temperate sublittoral to 30 m stenohaline 240 7–40 6–12 4
Western and sand covered
Australia rocky substratum
Hydrocharitaceae acoroides Indian Ocean, marine, protected tropical (25) Intertidal (in usually marine 26–35 3–150 12.5– 2–6
Enhalus Tropic. and exposed water, water filled away from 17.5
W.Pacific muddy sand and depressions) to freshwater
coarse coral rubble about 6 m inflow
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Genus Species Distribution Habitat Temperature (° Depth Salinity (%o) Leaf Leaf Leaf No. of
C) longevity length width leaves
(days)* (cm) (mm)
Thalassia hemprichii Indian Ocean, marine, sand and rocky tropical sublittoral stenohaline 40 10–40 4–11 2–6
Tropic. W. Pacific substratum, dead coral low water
reefs to 5 m
Thalassia testudinum Caribbean, Gulf of marine tropical (20–35) low water stenohaline (13.5– 24 10–60 4–12 2–6
Mexico to about 21)
10m
Halophila ovalis IndoWest. Pacific marine and estuarine tropical and mid tide to euryhaline (Shark 11–24 35– 1–4 ( 5–20 in pairs
(Swan and warm temperate 10–12 m Bay Swan Estuary 55, 7)
Canning Estuaries) (above 10) 25–35 optima, 10–
45 tolerated)
Potamogetonaceae
Ruppia maritima Cosmopolitan fresh, brackish, marine temperate and – euryhaline – 15–25 0.4–1 –
and hypersaline both subtropical
permanent and
ephemeral
Posidoniaceae
Posidonia oceanica Mediterranean exposed to moderately (17–20) sublittoral stenohaline 120 40–50 5–9 4–6
sheltered open shores to 30–40 m
and heads of bays
Posidonia angustifolia Southern Australia open nearshore waters temperate sublittoral stenohaline – (10) 4–6 2–3
rather than sheltered 2–35 m 20–75
embayments
Posidonia australis Southern Australia marine, sheltered temperate 0–14 m (35–55) 67–75 20–45 (6) 2–3
embayments and near (WA), 91– 10–15
coastal 125 (NSW) (20)
Posidonia sinuosa Southern Australia marine, sheltered temperate 1–35 m stenohaline 84–168 (2) (4) 7– 1–2
embayments and near 30–70 9 (11)
coastal areas with less (120)
sediment deposition than
P. australis.
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Genus Species Distribution Habitat Temperature (° Depth Salinity (%o) Leaf Leaf Leaf width No. of
C) longevity length (mm) leaves
(days)* (cm)
Zosteraceae marina N. Pacific, N. marine, bays Arctic to Tropic generally shallow marine, 55–83 120 2–12 2–5
Zostera (Zostera) Atlantic and estuaries of Cancer in the sublittoral, 0 to 4 m polyhaline (3– (narrow and
Atlantic Roscoff, France and 32) wide leafed
greater depth in the forms)
Pacific to 30 m
Zostera capricorni NE Australia marine, sand warm temperate low water to about 6 polyhaline – 7–50 2–5 2–5
(Zosterella) and New and mud m.
Zealand
Zostera mucronata Southern extremely temperate upper middle hypersaline in – 2–22 0.75–1.75 2–5
(Zosterella) Australia sheltered intertidal summer
shallow bays,
sand and mud
Zostera muelleri South eastern marine, temperate intertidal euryhaline – 5–30 0.75–2 2–5
(Zosterella) Australia protected bays
sand and mud
Heterozostera tasmanica Southern marine temperate upper sublittoral to 8 stenohaline 40–144 5–25 1–3 7–10 on
Australia and m erect
Chile lignified
stems
Phyllospadix torreyi N.America, marine exposed temperate to intertidal and subtidal stenohaline – 50–200 0.5–1.5 3
Pacific surf areas subtropical about 15m.
* Classification following Kuo & McComb (1989) based on den Hartog (1970). Data following den Hartog (1970) unless otherwise specified as follows: Habitat: Enhalus acoroides (Brouns
& Heijs 1986); Posidonia angustifolia (Cambridge & Kuo 1979). Salinity: Halophila ovalis Shark Bay (Walker 1989) and Swan Estuary (Hillman et al. 1989); Posidonia australis and
Amphibolis antarctica (Walker 1989). Leaf longevity: Enhalus acoroides (Brouns & Heijs 1986); Thalassia hemprichii (Heijs 1984 quoted from Borowitzka & Lethbridge (1989)
calculation); Thalassia testudinum (Zieman et al. 1979); Halophila ovalis (Hillman 1986); Posidonia australis (Silberstein et al. 1986); Posidonia oceanica (Ott 1980); Syringodium
filiforme (Zieman et al. 1979); Halodule wrightii (Morgan & Kitting 1984); Amphibolis antarctica (Walker 1985); Zostera marina (Jacobs & Noten 1980); Heterozostera tasmanica
(Bulthuis & Woelkerling 1983b). Number of leaves and leaf size: P. oceanica (Mazzella & Ott 1984); P. angustifolia P. australis, and P. sinuosa (Cambridge & Kuo 1979); Thalassodendron
pachyrhizum (Kirkman & Cook 1987).
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tion where seagrass was growing, leaf length, leaf shape, growth or turnover rate of leaves,and the spatial and temporal availability and distribution of epiphyte
propagules.
Seagrass shoots continuously produce new leaves, which appear between the older ones. New leaves grow from a basal meristem so that the oldest part of the leaf is
the tip. Old leaves are shed regularly, at a rate that varies with habitat (e.g. depth), season, and changes in growth rate (Ott 1980, McComb et al. 1981, Orth &
Moore 1986). Therefore, the time available for an epiphytic community to develop varies within a shoot, and seagrass leaves are a dynamic substratum for epiphytes
and sessile epifauna (Ballantine & Humm 1975, Heijs 1984, Borowitzka & Lethbridge 1989).
In the Borowitzka & Lethbridge (1989) morphological classification, the main meadow forming genera Posidonia, Zostera and Thalassia are all in group 3.
However, some of the differences between these groups could affect epiphyte colonization (Table 1). The leaves of Zostera marina (2–12mm) are generally narrower
than those of Thalassia (4–12mm), Posidonia australis (10–15mm) and P. sinuosa (4–11 mm). P. oceanica has four to six leaves per shoot (turion), whereas P.
australis has 2–3 leaves and P. sinuosa 1–2 leaves. Leafturnover times also vary, e.g. 65–75 days for P. australis, 84–108 days for P. sinuosa, 120 days for P.
oceanica and 5–83 days for Zostera marina (Table 1).
A group 4 species, Halophila ovalis (or “paddle weed”), has delicate, shortlived leaves (11–24 days). Although it is not a major meadow species, it is an
important colonizing species (Kirkman 1985) and has high productivity in some marine and estuarine communities (Hillman 1986). The other main meadowforming
species—Amphibolis antarctica and A. griffithii—are placed with Thalassodendron and Heterozostera in group 5. The genus Amphibolis is endemic to Australia.
Commonly called “sea nymph” or “wire weed”, this plant has a longlived, erect, lignified, cylindrical stem with small, shortlived (28–40 days; Walker 1985) leaves
arranged in terminal clusters. Amphibolis epiphytes have been studied by Kendrick (1983), Harlin et al. (1985), Kendrick et al. (1988), Borowitzka & Lethbridge
(1989) and Borowitzka et al. (1990). Heterozostera tasmanica, which also has an upright, lignified stem, is a much smaller species with fine, shortlived leaves and
seasonal growth. It is the main meadowforming species in the wide, shallow estuaries of the east coast of Australia (Howard 1982), but on the west coast of Australia
it is found in patches within Amphibolis and Posidonia meadows (Kirkman 1985).
In addition to differences in morphology, there are major differences in the habitats of the seagrasses. In the northern hemisphere, Zostera marina, the most
extensively studied temperate species, is found in predominantly shallow estuaries that are typically sheltered, lowenergy zones (Table 1). Posidonia oceanica from
the Mediterranean is more comparable with Western Australian Posidonia in morphology and habitat and is found to 40m depth (Table 1). P. sinuosa is found in
Western Australia to depths ranging between about 15m on the west coast (Cambridge & Kuo 1979) to up to 47m along the southern coast (C.Manning, CSIRO
Marine Labs, Marmion, pers. comm.) where the clear, sedimentfree water allows light to penetrate to greater depth.
The great diversity of seagrass species, morphologies and habitats offer very varied conditions for epiphyte colonization, survival and growth. In view of this
variability, we suggest caution when comparing epiphytegrazer interactions from different areas, particularly from different biogeographic provinces.
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Epiphytes
Numerous studies have described the epiphytic macro and microfloral assemblages associated with marine and estuarine vegetation (see Harlin 1980 for a review).
The term epiphyte is variously defined. Borowitzka & Lethbridge (1989), for example, used it to describe both the flora and fauna attached to the exterior surface of
plants and algae. This included sessile fauna, such as the Porifera, Cnidaria (Hydrozoa), Bryozoa, Polychaeta and Chordata (Ascidicea). We use it for a plant living on
the exterior of another plant (van Montfrans et al. 1984).
The other definitions we use are based mainly on those of van Montfrans et al. (1984) and are given below:
Epiphytes are collectively defined as both the microalgae and macroalgae colonizing the external surface of seagrass.
Periphyton are the microalgae (diatoms and microscopic filamentous forms), algal propagules, bacteria, microfauna and particulate material found in the mucuslike
layer that commonly coats seagrass leaves.
Detritus is usually described as finely abraded, or disintegrating material. However, it is also used for dead particles of seagrass and even small living algae (in which
sense it overlaps with the term periphyton) (Kitting 1984).
Microalgae include bluegreen algae and diatoms consisting of single cells that may be in loose chains but do not have holdfasts for attachment to the substratum.
Macroalgae include the erect, fleshy forms that colonize seagrasses. The macroalgae have been classified by their thallus morphology and life histories (Littler &
Littler 1980). Steneck & Watling (1982) modified the classification on the basis of the size and toughness of the plant and used it to group gastropod grazers by their
feeding characteristics.
Filamentous algae such as Cladophora and Ectocarpus consist of long chains of cells and attach by a holdfast.
Foliose algae such as Ulva and Rhodymenia have a thallus developed into a thin sheet, one or two cells thick.
Corticated macrophytes such as Acanthophora, Chondria and Bryothamnion have a larger, more complex thallus, several cells in thickness and with some
differentiation of cells and tissues.
Articulated calcareous algae such as Amphiroa, Corallina and Metagoniolithon are segmented, branched corallines whose branches consist of cells
impregnated with calcium carbonate, separated by uncalcified cells at the joints.
Encrusting algae with heavily calcified, prostrate growth over the substratum, include calcareous coralline species growing on seagrass leaves. Encrusting coralline
algae are among the most ubiquitous of seagrass epiphytes. They are pioneer colonizers, which develop rapidly to sexual maturity. Their low profile reduces the
likelihood of detachment from the host (Ducker et al. 1977, Bramwell & Woelkerling 1984, Borowitzka & Lethbridge 1989).
Drift algae are epiphytes that become detached from the seagrass but remain within the seagrass meadow (Virnstein & Carbonara 1985, Kendrick et al. 1990).
This term also refers
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to macroalgae originating outside the seagrass bed or attached to seagrasses by means other than holdfast (e.g. entanglement).
The floristic composition of the epiphytic algal community is determined by the occurrence of freeliving, reproductive stages of potential epiphytes, aspects of the
seagrass substratum (see p. 115) and abiotic environmental factors such as light, temperature, salinity, water movement, sediment scour, and nutrients that affect both
the seagrass host and epiphytes.
The epiphytic community is mainly composed of ephemeral species that can settle, grow and reproduce in a period dictated by the life of their host (Kendrick &
Hawkes 1988). Successful epiphytic species, therefore, have a more rapid turnover time than the seagrass substratum. Few epiphytes are thought to be obligate
epiphytes of particular seagrass species and most will grow on other substrata if conditions are favourable (Humm 1964, Harlin 1975, 1980, Bramwell & Woelkerling
1984, Orth et al. 1984, Borowitzka & Lethbridge 1989). However, many members of the Rhodophyta have only been documented as epiphytes (Kendrick et al.
1988, 1990).
Techniques for measuring epiphyte abundance

It is sometimes difficult to compare studies of the abundance of epiphytes because different methods have been used. The percentage cover of particular species on the
seagrass surface has been estimated, and classified in groups following a modification of the BraunBlanquet method (Van der Ben 1969, Ballantine 1979, Jacobs et al.
1983). Cell counts have been related to surface area or volume scraped from the seagrass surface (Neckles et al. 1994). The density of crustose corallines on leaves
has been measured (Harlin et al. 1985). The biomass of epiphytes has been measured and quoted as fresh weight, ashfree dry weight and as a percentage of leaf
biomass (Bulthuis & Woelkerling 1983a, Heijs 1984, Silberstein et al. 1986, Neckles et al. 1993). The chlorophyll a content of epiphytes scraped from the seagrass
surface has been used as an index of biomass (Borum 1987). Oxygen production and carbonate as 14C uptake have also been measured to estimate epiphyte
abundance (Penhale 1977, Mazzella & Alberte 1986).
Interactions between epiphytes and seagrasses

Seagrasses positively affect epiphytes by providing a settlement surface and possibly nutrient exchange (Harlin 1975, 1980, Walker & Woelkerling 1988). Early
studies suggested that seagrasses were “leaky” and that carbon and nutrient exchange might enhance epiphytic growth (McRoy et al. 1972, McRoy & Goering 1974).
However, more recent studies have shown that earlier estimates of nutrient release from seagrass leaves may be excessively high due to experimental artefacts (Brix &
Lyngby 1985). These authors suggested that the main source of phosphorus and nitrogen for epiphytes must be the surrounding water and not the seagrass, which
releases nutrients too slowly to support the epiphyte growth observed in the field. The current view is that most epiphytes utilize the seagrass host only as a physical
substratum for settlement and do not derive nutrients direct from the host (Orth & van Montfrans 1984). This view is supported by the similarity between epiphytes
found on artificial substrata and those recorded from seagrasses of similar morphology (Virnstein & Curran 1986, Horner 1987, Hall & Bell 1988, Lethbridge et al.
1988). However, Orth et al. (1984) consider that there may be significant nutrient exchange with the microalgal component of the flora that is more intimately
associated with the seagrass surface. For example,
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bacterial populations on Zostera marina may be supported almost entirely by carbon obtained directly from seagrass photosynthesis (Kirchman et al. 1984).
Conversely, soluble nitrate excreted by bluegreen algae on Thalassia testudinum may be used by the seagrass (Goering & Parker 1972).
Seagrasses may deleteriously affect epiphytes by inhibiting their growth through the release of chemical compounds such as phenolics. Phenolic compounds have
been recorded from a number of seagrasses (Zapata & McMillan 1979) and chemical extracts from Zostera marina have an inhibitory effect on microalgae and
bacteria (Harrison & Chan 1980, Harrison & Durance 1989). However, extracts from Posidonia australis, Halophila ovalis and Amphibolis antarctica showed
almost no toxic or inhibitory effects on a range of microorganisms and there appears to be little evidence that seagrasses generally inhibit the growth of algal epiphytes
(Reichelt & Borowitzka 1984).
The effects of epiphytes on seagrasses may be either beneficial or detrimental (SandJensen et al. 1985, Silberstein et al. 1986, Orth 1992). Probably the most
severe negative effect of epiphyte growth is that of shading, as has been observed in meadows of Heterozostera tasmanica (Bulthuis & Woelkerling 1983a),
Posidonia australis (Cambridge et al. 1986, Silberstein et al. 1986) and Zostera marina (SandJensen 1977, SandJensen & Borum 1983, Borum 1987).
Attenuation of light limits the depths at which marine macrophytes grow (Cambridge & McComb 1984, Masini et al. 1990). Thus, a reduction in photosynthetic light
due to epiphytic growth may decrease the depth at which seagrasses can grow (Cambridge and McComb 1984, Silberstein et al. 1986, Shepherd et al. 1989, Masini
et al. 1990).
Epiphytes create a boundary layer around the seagrass leaf through which nutrients and oxygen must diffuse to be available for the seagrass’s metabolism. This
unstirred layer can interfere with the uptake of carbon and phosphorus in Zostera marina (SandJensen 1977, SandJensen et al. 1985) and Enhalus (Johnstone
1979). Epiphytic growth may also be selflimiting through similar light attenuation and boundary layer effects (Borum 1987). SandJensen et al. (1985) placed
microprobes within the epiphyte layer and found that the photosynthetic activity of epiphytes contributed to unfavourable O2:CO2 ratios for seagrass and macroalgal
photosynthesis and produced anoxic conditions at the leaf surface. However, these experiments were conducted in a shallow estuary; in higher waveenergy
environments, boundary layer effects of epiphytes inhibiting the passage of oxygen and carbon dioxide may be less.
The detrimental effects of epiphytes on seagrass photosynthesis is not proven (Jacobs & Noten 1980, Hootsmans & Vermaat 1985, Mazzella & Alberte 1986,
Borum 1987). Jacobs & Noten (1980) suggested the lack of effect on Zostera marina was because the seagrass adapted to the epiphyte community throughout the
year. Similarly, Mazzella & Alberte (1986) found that Zostera marina with epiphytes photosynthesises at lower light intensities than those usually measured within the
area. They also point out that the epiphytic community of diatoms and red algae characteristically absorb the green portion of the light spectrum, which is essentially
unused by Zostera marina.
Epiphytes may also increase the susceptibility of seagrasses to leaf loss when subjected to physical forces such as waveaction. In zosteroid seagrasses (e.g.
Posidonia and Thalassia), heavy epiphyte loads may reduce leaf flexibility, leading to loss of older sections of leaf (Jacobs et al. 1983, van Montfrans et al. 1984,
Heijs 1985a,c). In amphiboloid types (e.g. Amphibolis), a high biomass and large size of epiphytes may increase drag on the seagrass during wave surges, such that
leaves and stems may be torn off and whole plants ripped from the substratum (Borowitzka & Lethbridge 1989).
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The positive effects of epiphytes on seagrass have been discussed less often than the negative effects. If epiphyte production exceeds that of the seagrass host,
detrital material contributed to the system could, on breaking down, be available for seagrass nutrition (Penhale & Smith 1977, Mazzella & Alberte 1986). Epiphytes
may reduce water movement (Borowitzka et al. 1990) and protect littoral seagrasses at low tide from desiccation (Penhale & Smith 1977) and excess insolation
(Trocine et al. 1981).
Epiphyte productivity may also contribute significantly to the overall productivity of seagrass meadows. Over 60% of the maximum photosynthetic rate of Posidonia
australis in Cockburn Sound, Western Australia, was attributed to epiphyte photosynthesis (Silberstein et al. 1986). Epiphytes on Zoster a marina in Massachusetts
accounted for 27–50% of the total photosynthetic rate (Mazzella & Alberte 1986). Epiphytes on Amphibolis spp. in Western Australia had a similar biomass to the
seagrass host in spring, the period of maximum biomass (Edgar 1990).
In summary, there is a great diversity in the species, morphologies and life histories of epiphytes. The timing and intensity of their recruitment, local physical conditions
such as water temperature and wave action, the morphology and life history characteristics of the seagrass substratum, and the availability of space on the seagrass leaf
may all influence the outcome of the interaction of epiphytes and seagrasses. Another factor affecting this interaction is the influence of grazers.
Grazers
Grazing is an important process within seagrass ecosystems (Howard 1982, Orth & van Montfrans 1984, Valentine & Heck 1991, Mazzella et al. 1992 and many
others) for three main reasons: (a) it provides a major trophic link for the cycling of nutrients (Robertson & Mann 1980, 1982, van Montfrans et al. 1984, Klumpp et
al. 1992), (b) it affects the productivity of the seagrass communities (van Montfrans et al. 1982, Wetzel & Neckles 1986, Orth 1992, Neckles et al. 1993) and (c) it
can change the species composition of seagrass epi4 phytes (van Montfrans et al. 1982, Jacobs et al. 1983).
The diversity of grazers within seagrass meadows is large (van Montfrans et al. 1984, Klumpp et al. 1989, Lanyon et al. 1989, Orth 1992). Grazers can be
described in terms of their taxonomic grouping (e.g. fishes, molluscs, crustaceans; Klumpp & Nichols 1983, Pollard 1984, Nielsen & Lethbridge 1989); feeding
strategy/behaviour (e.g. generalist browsers, sweepers, rakers, shovellers; sensu Steneck & Watling 1982); size of food ingested (macrophagous, microphagous;
Zimmerman et al. 1979) or size of grazer. Small invertebrate grazers larger than an average copepod but smaller than about 2.5cm are termed mesograzers by Brawley
(1992). They include small crabs, shrimp, amphipods, isopods, dipteran larvae, small molluscs and polychaetes. Some animals grow through a “mesograzer” stage. The
term micrograzer was originally proposed for this group of organisms; however, it is now restricted to much smaller (less than 2mm) herbivores, such as flagellates
(Neckles et al. 1994). The 2mm limit is arbitrary, but is reported to be the smallest size at which most organisms can be detected underwater by the unaided eye
(Brostoff 1988).
Sea urchins are the only major taxa that graze directly on temperate seagrasses (Mazzella et al. 1992) but many other taxa graze on epiphytes, either directly or
indirectly. Direct grazing may be selective, such as cropping the diatom crust close to the surface of the seagrass blades, e.g. radular grazing by small gastropods (van
Montfrans et al. 1982, Klumpp et al. 1992), or plucking small filamentous macroepiphytes from the periphyton layer, e.g. by
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shrimps using chemosensory chelae (Morgan 1980, Howard 1982). Direct grazing may also be nonselective as when freeliving amphipods remove the loosely
adhering layer of diatoms and flocculent detritus from seagrass leaves (Zimmerman et al. 1979, Howard 1982). Indirect grazing is the ingestion of plant material during
feeding on associated fauna (Klumpp et al. 1989).
Another grazer feeding mode is filter or suspensionfeeding on epiphyte propagules. Filter feeders are found in a number of faunal groups including hydroids,
bryozoans, bivalve molluscs, holothurians and ascidians (sea squirts). Some species of hydroids, bryozoans and ascidians may be attached to leaves or erect stems of,
for example, Amphibolis and Thalassodendron (Borowitzka et al. 1990, Hughes et al. 1991). Others, such as the molluscs, holothurians and ascidians, are found on
the sediment.
We will describe the major grazers in seagrass meadows by their taxonomic groupings and, where appropriate, address their modes of feeding.
Types of grazers
Fishes
Direct herbivory by fishes on seagrasses is rare (Bell & Pollard 1989) except in the hemiramphids (Robertson & Klumpp 1983) and monacanthids (Bell et al. 1978,
Klumpp & Nichols 1983). However, mugilids, hemiramphids and monacanthids as well as some kyphosids and sparids have been reported to eat epiphytic algae in
seagrass meadows. Most of these fishes appear to be feeding mainly on the associated epifauna (Klumpp et al. 1989, Mazzella et al. 1992).
In a review of trophodynamics in seagrasses, Klumpp et al. (1989) suggest that direct grazing of Australian seagrasses by fishes is probably insignificant in terms of
seagrass removal. There is evidence that the ingestion of seagrass may be incidental, with the preferred diet being epifauna and epiphytes living on the seagrass leaves
(Bell et al. 1978). But there is also some evidence that monacanthids and hemiramphids gain some nutrition from the plant material (Conacher et al. 1979, Klumpp &
Nichols 1983).
Echinoderms
Sea urchins are primarily generalist browsers that eat mainly plant material and a variety of sessile encrusting animals but can exhibit marked dietary preferences when
food resources are freely available (Hawkins & Hartnoll 1983). They are capable of consuming large areas of seagrass directly (Lawrence 1975, Greenway 1976,
Mazzella et al. 1992, Klumpp et al. 1993), particularly when epiphyte biomass is low, or sea urchin numbers are high (Cambridge 1980, Ogden 1980, Larkum &
West 1990). Klumpp et al. (1993) estimated that sea urchins grazed about 24% of the annual aboveground leaf production of Thalassia hemprichii meadows in the
Philippines, although this impact may vary from less than 5% to 100% at different times of the year depending on sea urchin density and population structure. Sea
urchins have been reported to feed on the epiphytized tips of seagrass leaves and on drift algae (Ott 1980, Kirkman & Young 1981, Mazzella et al. 1992). Klumpp et
al. (1993) found that the sea urchin Tripneustes gratilla consumed Thalassia hemprichii fronds in preference to detached seagrass debris or the red alga Amphiroa
fragilissima, whereas the sympatric sea urchin Salmacis sphaeroides exhibited no preferences and consumed all three food types in proportions that varied with their
availability. Thus, some sea urchins have the ability to change diet depending upon local conditions.
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In temperate waters, sea urchins may have caused localized but severe damage to seagrass meadows but it is not clear whether they were responsible for the
seagrass decline or were a secondary effect on meadows under stress. For example, Cambridge et al. (1986) suggested that the sea urchin Temnopleurus
michaelseni was responsible for the loss of seagrass in Cockburn Sound (Western Australia). However, as heavy densities of sea urchins were present in the Sound
years before the seagrasses declined, and although they cleared large areas of deteriorating seagrass meadows, they are not thought to be the primary cause of the
meadows’ decline (Shepherd et al. 1989).
Asteroids feed on a variety of material and, while many species are carnivores, some species or stages in the life cycle may be herbivores (Hawkins & Hartnoll
1983). Most species are not selective feeders. During feeding, the stomach is extended over the substratum, and microalgae, macrofauna and detritus are absorbed
(Hawkins & Hartnoll 1983). They are probably not important grazers of seagrasses or epiphytes and there have been no published accounts of significant effects.
Molluscs
Molluscs are one of the main feeders on seagrass epiphytes. Grazing by prosobranch snails has been well documented (Lubchenco & Gaines 1981), but other
gastropods such as seahares (Aplysia spp.), which often live in seagrass beds, also eat epiphytes (Kikuchi & Pérès 1977).
Prosobranch snails are generalist grazers of seagrass epiphytes, although their ability to remove different sizes, morphologies, and therefore species, of algae is
influenced by their method of feeding. Different species of molluscs have different types of radulae with different cutting styles and abilities to remove plant material.
Steneck & Watling (1982) considered that dietary differences among herbivorous molluscs were related to the design of their radula and the morphology of algae. In
their model, prosobranch gastropods and chitons are divided into four groups based on the form of the radula. The algae are divided into seven groups based on
morphology, size and toughness (following an earlier scheme by Littler & Littler 1980). The model predicts that the smallest algae with simple morphology (e.g. the
periphyton that first colonizes seagrass leaves) will be grazed most readily, with grazing difficulty increasing with algal size, toughness and complexity of structure. In
increasing order of difficulty the microalgae will be followed by filamentous forms, foliose forms, corticated macrophytes, leathery macrophytes, articulated calcareous
algae and crustose coralline algae. Peduzzi’s (1987) results support this model. He found that herbivorous gastropods preferred small simple algae (diatoms and green
algal films) to macroepiphytes and seagrass detritus.
Steneck & Watling’s (1982) functional model has been widely quoted in the literature. However, application of the model to all grazing types, particularly to grazing
on seagrass epiphytes, may be simplistic. Padilla (1985) found that, contrary to the model, calcareous algae were often grazed, and she suggested that they were brittle
and easily removed. Similarly, Nielsen & Lethbridge (1989), working on epiphytes of Amphibolis griffithii in Western Australia, found a large proportion of
calcareous algae in the diet of the trochids Thalotia conica, Prothalotia lehmanni and the columbellid Pyrene bidentata. This pattern, however, may also be
because of differential digestion times, with coralline algae taking longer to digest.
Filterfeeding molluscs
Filterfeeding molluscs are also found in seagrass beds (Irlandi & Peterson 1991). They are usually considered to be herbivores, filtering phytoplankton as food (Bayne
& Hawkins
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1992). Their role in grazing propagules of epiphytes is unknown, but is most likely insignificant compared with the grazing activities of gastropods.
Crustaceans
Crustaceans eat macrophytes, microphytes and debris associated with the seagrass substratum (Kirkman 1978, Zimmerman et al. 1979, Morgan 1980, Howard 1982,
Morgan & Kitting 1984). Hawkins & Hartnoll (1983), however, comment that many are omnivorous, and even among strictly herbivorous species, few special
adaptations of the grazing mechanism have been observed. Hence these species may have the ability to switch among food types when one type becomes scarce.
Crustaceans that may be significant grazers of seagrass epiphytes include copepods, amphipods and isopods. Harpacticoid copepods are abundant on seagrasses
and macroalgae. They feed predominantly on the periphyton and may in turn be a major dietary component of fishes in the seagrass habitat (Hicks 1986).
Amphipods have a variety of feeding behaviours, eating larger macroalgae and grazing on microscopic algae in the periphyton and detritus (Zimmerman et al. 1979).
Many species can change their feeding method to utilize different foods that become available (Klumpp et al. 1989). Howard (1982) observed that Paradexamine
churinga in southern Australia usually fed by antennal filtering, but if epiphytic diatoms were abundant and large enough, it used the gnathopods to collect these from
the seagrass surface. The effects of amphipod grazing may vary because populations of amphipods fluctuate with environmental conditions, fish predation and food
availability (e.g. Young & Young 1978, Stoner 1980a,b, 1983, Edgar 1990, Gambi et al. 1992).
Isopods can be extremely abundant in seagrass meadows (Klumpp et al. 1989). Some forms are specialized for life in the habitat, with cryptic colouring and
modifications to the body that aid concealment or attachment to the elongate seagrass. Little is known of the diet of individual species. Some isopods consume algae
and seagrass tissue (Nienhuis & van Ierland 1978, Hootsmans & Vermaat 1985, Brearley & Walker 1993), biting off small portions with the mandibles, and using
other mouthparts to hold the food and push it into the mouth. In the Netherlands the idoteiid isopod Idotea chelipes was the main consumer of the seagrass Zostera
marina, although it accounted for less than 3% of the total annual seagrass production (Nienhuis & van Ierland 1978). In Western Australian Posidonia meadows, leaf
boring isopods (Limnoriidae: Lynseia) have been found living within the leaf and consuming mesophyll tissue (Brearley & Walker 1993). Several species have been
reported to graze on epiphytes (Nicotri 1980, Hootsmans & Vermaat 1985, Borum 1987, Williams & Ruckelshaus 1993).
Decapods are also found in seagrass meadows. Many species are carnivorous (Bell & Westoby 1987) but those that are herbivorous predominantly feed on
seagrass tissue (Klumpp et al. 1989).
Food selection and feeding patterns

The factors influencing food selection are complex and may be a function of the feeding apparatus, or it may be such features as palatability, availability, energy content,
ease of digestion or absorption efficiency that determine the choice of a particular food. Nicotri (1980), who looked at the food preferences of intertidal amphipods,
suggested that food preferences are not directly determined by the nutritional quality of the algae, but are significantly influenced by the morphological and
physiognomical characteristics of the plant relative to the animals. Epiphytes are generally regarded as more attractive than the
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seagrasses as food for grazing. They have a high biomass and higher nutritive value with different structural storage material (Nicotri 1980, Klumpp et al. 1989).
Mesograzers probably have a central role as controllers of algal periphyton and as vehicles for energy transfer from epiphytes to higher consumer levels (Gambi et al.
1992, Mazzella et al. 1992). The distribution of preferred food items may explain the microdistribution patterns of Baltic idoteiids (Salemaa 1987) and of trochid
molluscs on Posidonia oceanica in the Mediterranean (Mazzella & Russo 1989).
Little is known of the biology and dietary preferences of many grazers, particularly the smaller species. This may be due in part to their cryptic habits during daylight,
when most sampling is done (Kitting et al. 1984). In a study conducted at night, with remote camera imaging, these authors found two species of shrimp, previously
considered to be detrital feeders, feeding on seagrass epiphytes. Conflicting reports in the literature on the diet of some species may be because of variations in
abundance of preferred foods, seasonal conditions and the age of the grazer, as well as different methods of investigating diet (Kitting 1984). Edgar (1992) suggests
that guilds of mobile amphipods move among seagrass meadows on the southern Australian coast, in response to available food. Although Edgar (1992) was
principally referring to amphipods feeding on detritus and detached macrophytes, these species are also known to feed on seagrass epiphytes. Grazers may also use a
variety of algae as food and habitat. Buschmann (1990) reports that low numbers of amphipods were collected on the preferred food source during daylight; most
amphipods were found within intertidal turfs and migrated at night to feed on the preferred food.
Measurements of grazer diets

The methods used to determine the components of the diet of small grazers include examination of grazing trails, gut contents and faecal pellets with light and scanning
electron microscopy (van Montfrans et al. 1982). However, there may be a problem with determining diet by visual analysis of gut contents (Kitting 1984), since most
material in the gut resembles detritus, regardless of origin. This may have resulted in grazers being assigned to detritivore status. Diet has also been studied by observing
feeding behaviour (Howard 1982, Robertson & Mann 1982), analysis of body tissues isotopically and comparing them with potential food sources (Fry 1984, Kitting
et al. 1984) and food choice experiments (Zimmerman et al. 1979, Nicotri 1980). These experiments can sometimes be difficult to interpret and extrapolate to natural
conditions because the range of food offered does not always reflect the wide variety of food available in the field.
Removal of algal biomass by grazers has also been demonstrated experimentally (Kirkman 1978, Howard 1982, van Montfrans et al. 1984) (Table 2). In the field,
cages and containers have been used to exclude grazers to compare the epiphytes on grazed and ungrazed seagrass plants (Robertson & Mann 1982, Klumpp et al.
1992, Jernakoff & Nielsen in press). Experimental studies of grazerepiphyte interactions have also been undertaken in aquaria and mesocosms (Caine 1980, Brawley
& Adey 1981, Howard 1982, van Montfrans et al. 1982, Neckles et al. 1993). Some studies of epiphyte grazers have been limited to descriptions of the different
appearance of the seagrass surface before and after grazer activity.
In summary, grazers of seagrass epiphytes show a wide diversity of shapes, sizes, taxa, species and feeding morphologies. They occupy a range of feeding niches in
seagrass meadows. Many appear to be generalist feeders, although little is known about their dietary preferences. Grazers can potentially have a significant impact on
epiphyte abundance in seagrass meadows, but the nature of this impact may vary in space and time.
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Table 2 Summary of selected studies of grazers of seagrass epiphytes (table based on Table 2 of van Montfrans et al. 1984). (A)=amphipod; (G)
=gastropod; (C)=carid; (D)=decapod; (I)=isopod; (U)=sea urchin.
Grazer Seagrass, location, study Description of study/results Reference Comments on experimental design
type
Modulus Thalassia testudinum, Build up of fouling organisms on tiles was considerably less in areas of Mook 1977 Uncaged treatments were assumed
modulus (G) Halodule wrightii. seagrass, compared with areas of bare sand. Within sand patches, to be grazed although there are no
Indian River Lagoon, Ft caged tiles without grazers had more fouling organisms than uncaged observational data on the presence
Pierce, Florida, USA. tiles, However, there was no difference in the amount of fouling on of grazers in these treatments.
Field and lab. experiments, caged and uncaged tiles within seagrass beds. Significantly more snails
collections from subtidal were found on tiles within seagrass beds and there was a negative
meadows, depth not correlation between snails and the amount of fouling. A separate
specified experiment demonstrated that snails could reduce the amount of
fouling.
Cymadusa Halodule wrightii, Feeding behaviour and dietary preferences for particular algae were Zimmerman Algae used in feeding trials were
compta (A) Syringodiumfiliforme, demonstrated (see also Table 3 for rates). C. compta fed et al. 1979 offered separately and thus the
Gammarus Ruppia maritima, preferentially on macroepiphytes, drift algae and detritus. G. results of feeding preference
mucronatus (A) Thalassia testudinum. mucronatus fed equally on macroepiphytes and detritus. M. nitida experiments may be due to
Melita nitida Indian River, Lagoon, Ft. mostly consumed macroepiphytes. G. bonnieroides fed mostly on laboratory artefacts because the
(A) Pierce, Florida, USA. Less microepiphytes (diatoms) and detritus by grooming the leaf surface. trials did not examine the range of
Grandidierella than 2 m 14
C studies showed epiphytic algae provided 41–75% of food ingested. epiphyte species concurrently in the
bonnieroides 14 field.
(A) Drift algae varied between 11 and 24% of C uptake by amphipods.
There may be some resource partitioning within amphipod community.
However, predation and an overabundance of food may indicate that
evidence for resource partitioning may be an artefact.
Caprella Zostera marina. In the absence of this caprellid amphipod, periphyton production on Z. Caine 1980 Densities of 177 individual grazers
laeviuscula. (A) Washington Sound USA. marina leaves increased by 411%. It is suggested that the presence of per leaf cluster were reported. No
Field and laboratory studies; this amphipod may allow eelgrass to grow in areas where it would data were given on epiphyte
collections from littoral zone otherwise be excluded due to competition with epiphytes for light. biomass. There is no clear
statement as to method of
assessing periphyton removal or if
grazing rate was based on
periphyton on Z. marina leaves or
on glass slides, which were both
placed in aquaria.
Palaemonetes Halodule wrightii. P. pugio fed heavily on epiphytes, but did not consume seagrass. Morgan Two shrimps were used in each
pugio (C) Corpus Christi, Texas. Large shrimps also consumed mysids at a rate of 6 per day, but 1980 experiment with 3–5 replicates.
Experiments were repeated a
number of times with
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Grazer Seagrass, Description of study / results Reference Comments on experimental design

location, study
type
did not appear to search for prey. Rather, prey was fresh seagrass and epiphytes. 14C studies to estimate
consumed when encountered. Dead mysids were also food consumed,
consumed. Epiphyte material was a heterogeneous mix of
filamentous and single celled diatoms, bluegreen algae,
microscopic animals, fine particulate material and bacteria.
Grazing rate increased with temperature (14–30°C), but not
with salinity (11–33‰).
Assimilation efficiency average 83% and was not affected
by temperature or salinity. Shrimps 6–12 mg (11–15 mm
total length) used in experiments, for shrimps to grow from
11–24 mm, assuming a 10% growth efficiency, 6.1 mg DW
of epiphytes must be consumed daily. See also Table 3 for
rates.
Paradexamine Heterozostera Amphipods removed epiphytes. In a chamber without Howard Experiment not replicated. Densities of grazers in
churinga (A) tasmanica. epiphytes, detrital material accumulated on leaves. The 1982 aquaria similar to that found in Z. muelleri beds found in
Tethygeneia Western Port Bay, appearance of ungrazed and grazed leaves are described. the same areas as H. tasmanica (P. churinga 120
nalgo (A) Victoria, Australia. adults (=1430 m−2) and T. nalgo 65 adults and juveniles
Aquaria
experiment and (770 m−2). P. churinga was a suspension feeder, with
field observations only detrital material in the gut. T. nalgo consumed fine
plant material.
Littorina Zostera marina. L. neglecta consumed epiphytes (filamentous algae and Robertson Field experiment used cages to exclude grazers;
neglecta (G) Chezzetcook Inlet, diatoms) as well as detrital material from the surface of & Mann experiment ran for 26 days. Removal of epiphytes by
Halifax, Nova leaves. Exclusion experiments showed epiphyte populations 1982 grazers is discussed in respect to enhanced growth and
Scotia. were reduced by these grazing snails, and it was suggested survival of eelgrass as well as importance in nutrient
Field experiment, that there was a limited food supply. In treatments without cycling (however van Montfrans et al. 1984, review
0.75 m depth snails growth of diatoms and the settlement of detritus implies that this was demonstrated rather than
increased. discussed).
Bittium varium Zostera marina. Grazing removed periphyton biomass on artificial seagrass van The authors suggest that differences observed between
(G) Chesapeake Bay, by 62.7%. Grazing appeared to be selective; senescent Montfrans the periphyton on artificial and living Z. marina may
Virginia, USA. diatoms constituted bulk of the periphyton and were et al. 1982 need to be considered before more studies are
Aquaria selectively removed and Nitzschia sp. and Amphora sp. undertaken, although they consider that grazing on
experiments were grazed before more tightly adhering Cocconeis periphyton was still effectively demonstrated by the
scutellum. See also Table 3 for grazing rates. short term SEM study.
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Grazer Seagrass, Description of study/results Reference Comments on experimental design

location, study
type
Artificial seagrass colonized in field and used in laboratory
studies with snails. Chlorophyll a and phaeophytin and
periphyton biomass was measured.
Anachis avara Thalassia Documents gradients of epiphytes along leaf blades. Kitting 1984 Review article coupled with observation of feeding
semiplicata (G) testudinum near Feeding occurred most frequently amongst epiphytic algae behaviour. Observations made in the field using remote
Bittium varium Halodule wrightii at night, not during the day when gastropods were located photographic sampling coupled with microacoustic
(G) and Ruppia on the sediment. Suggests feeding on epiphytic algae may monitoring and immediate, high resolution gut analysis
maritima. be more common than literature reports suggest, many enabling comparison of foraging frequencies with
Gulf passages, animals may not be detritivores and may actively select availability of foods. Anachis 200 m−2≈1 per plant and
Texas. 1–2 m particular types of living algae. Kitting draws analogy to
depth forests and the importance of small ephemeral species, Bittium 2000 m−2 ≈10 per blade. No grazing rates given.
rather than the longer lived more massive species, in Has a table with gut contents predominantly of diatoms,
sustaining food webs. yet comments on feeding predominantly on the lightly
calcified coralline alga Heteroderma lejolisii.
Palaemonetes Halodule Epiphytic algae accounted for nearly half above ground Morgan & Productivity of seagrass and epiphytes was measured by
pugio (D) wrightii. biomass and primary productivity. They were also a major Kitting 1984 14C incubations. Remote controlled cameras were used to
Bittium varium Corpus Christi recognizable component in the diet of common seagrass record feeding behaviour. Diet was determined by gut
(G) Bay Southern associated invertebrates (table given of available foods and content analysis.
Anachis avara Texas. relationship to diet). Seagrass tissue was not included in the
(G) Hypersaline diet. Isopods were unusual in containing coralline algae and
Tozeuma (60‰) condition grass shrimps P. pugio sometimes consumed foramniferans
carolinensis in summer, very with the epiphytes. Shading of seagrasses by epiphytes was
(D) shallow location not a problem encountered by these seagrass populations
Penaeus 10–75 cm depth) and a hurricane did not appear to damage the meadows.
duorarum (D)
Cymadusa sp.
(A)
Callinectes sp.
(D)
Hydrobia Zostera marina. All grazer species caused a reduction in periphyton and Hootsmans Field observations showed that eelgrass was virtually free
ulvae (G) Eastern Scheldt, there was a corresponding increase in seagrass biomass. & Vermaat of periphyton during the growing season. This was
Littorina SW Netherlands. Idotea had the largest impact on periphyton, Littorina the 1985 probably due to large number of H. ulvae (up to 60000
littorea (G) Aquaria least. However, Idotea commenced grazing on seagrass individuals m−2 of bottom surface), Experiments were
Idotea chelipes experiment with when periphyton was removed. Seagrass growth was designed to describe effects of snails on eelgrass.
(I) collections from similar under low grazing intensity by Littorina and more Densities of grazers approximated field densities
intertidal seagrass substantial removal of periphyton by Idotea. As light (Hydrobia=100; Littorina=5 and Idotea=1) per tank.
bed intensity used in the experiment was below saturation level, Speculation to
the more substantial
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location, study
type
removal of periphyton by Idotea should have produced a greater account for no difference in growth between
increase in growth. It is suggested that Littorina may graze on the most grazer treatments despite differences in
active photosynthetic part of the leaves, compensating for the lower rate periphyton removal. Light at below saturation
of periphyton removal. See also Table 3 for grazing rates. levels,
Modulus Halodule Treatment without epifaunal grazers had higher biomass epiphytes and Howard & Cores of sediment and seagrass were
modulus (G) wrightii. higher leaf loss. Aboveground and belowground biomass and shoot Short 1986 transplanted in pots into large outdoor culture
Cerithium Microcosm density was lower where grazers were absent. tanks. Densities of grazers were high but
muscarum (G) expt, Florida realistic (2000) per pot. However, many
Bittium varium individuals left the pots and were found on the
(G) walls of the tanks. At the end of the experiment
Astyris lunata 155 individuals remained in the treatment pots.
(G)
Bulla striata (G)
Prunum
apicinum (G)
Cymadusa
compta (A)
Gammarus
mucronatus (A)
Grandidierella
bonnieroides
(A)
Erichsonella
attenuata (I)
Hippolyte
pleuracanthus
(C)
Idotea baltica Zostera Epiphyte accumulation was found in eutrophic conditions. Epiphytes Borum Used chlorophyll a as an index of biomass. Rate
(I) marina. increased exponentially with leaf age (10–14 days). Algal doubling time 1987 of periphyton removal in In units of chlorophyll.
I. viridis (I) Roskilde Fjord, was 3–5 days on young leaves increasing in older leaves to 14 days. The relationship of chlorophyll to biomass of
Sphareoma Denmark. Invertebrate grazing reduced algal doubling time to 5.8 days in older periphyton was not given.
rugicuda (I) Field leaves. Grazing accounted for bulk of epiphyte biomass loss. Turnover of
Theoduxus experiment: biomass by grazers accounts for increase in productivity to 4–5 times
fluviatilis (I) salinity 8–12‰ that which is indicated by biomass. If grazers were excluded and all
Littorina and 0.6–0.8 m other factors held constant, epiphyte biomass on older leaves would be
saxatilis (G) depth 10 times higher than actually found in the eelgrass bed. See also Table 3
Hydrobia sp. for grazing rates.
(G)
Gibbula Posidonia Dietary preference of trochids was compared using different Peduzzi Consumption of diatoms, algal films
umbilicarlis (G) oceanica. components of epiphyte populations and algal films grown on glass 1987 (Cladophora sp.) on glass slides, seagrass
Jububinus Gulf of Naples. slides. Comparison made between gut contents and faeces of debris and macrophytes on epiphytised leaves
striatus (G) collection from experimental and field collected snails. Consumption, absorption was examined. Organic carbon content of algae
2–3 m depth efficiency greatest when feeding on algal films. See also Table 3 for (≈23–25%). Faecal pellet and gut content
grazing rates. analysis, absorption efficiency was calculated.
Rates of consumption as per animal total ww
(living animal including shell). Conversion to
DW from Peduzzi 1987 (not seen).
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location, study
type
Gibbula Posidonia Documentation of epiphyte community along leaf axis (age). Mazzella & Descriptive study, with many photographs of epiphytes
ardens (G) oceanica. Youngest basal section colonized by bacteria and diatoms, ageing Russo 1989 and grazing marks,
G. Gulf of Naples. tissues by encrusting soft and calcareous algae, oldest section
umbilicarlis collection from (leaf tips) with overlying upright layer. Diet of snails determined
(G) 1m depth by observation of grazing trails and faecal pellets. Distribution of
two gastropod species along leaf corresponded to different dietary
preference. G. ardens fed diatoms in youngest section, G.
umbilicarlis preferred macroalgae in older section. Distribution of
snails also followed spatiotemporal distribution of epiphytes within
meadow.
Thalotia Amphibolis Snails reside within leaf clusters. Epiphytes were a more Nielsen & Descriptive study focusing on the diets of gastropod
conica (G) griffithii. important component of the snail diets than detritus. Trochid Lethbridge snails. Consumption rates are not given. Seasonality of
Prothalotia Penguin Island, gastropods with rhipidoglossan radulae consumed diatoms and 1989 some epiphyte species is mentioned, and a change in the
(G) Warnbro Sound, small encrusting coralline algae on leaves. Pyrene with a diet with season is noted. Abundance of individual
lehmanni Western rachiglossan radula consumed similar food items, but also species is contained in a table comparing summer and
(G) Australia. 3–4 m consumed larger items such as hydroids. Snail diets closely winter flora, However, changes with or without grazers
Pyrene depth reflected range and abundance of epiphytes found on leaves in are not given,
bidentata summer, during winter some diatom species were
(G) disproportionately present in diets. This may be due to selection,
or the patchy distribution of diatoms.
Strombus Enhalus Documents the production and distribution of epiphytes and Klumpp et The study was carried out over 4 months (February to
mutabilis (G) acoroides. grazing rates in tropical seagrasses. Some grazers migrated from al. 1992 May 1991). Samples were collected by selecting a pre
Cerithium Balinao, sediment to canopy to feed at night while others remain within the determined number of seagrass stems. No statistical
tenellum (G) Philippines. canopy and feed regardless of the time of day. Diet of grazing justification of the sampling method including the
Subtidal, snails showed no preference within periphyton. Coralline algae number of stems chosen was given. The study focused
sheltered were not grazed. The study uses a combination of laboratory and on grazing gastropods although the authors
nearshore sites. field studies with real and artificial seagrass. The study is set at acknowledge that grazing by crustacean grazers (and
nine sites but is concentrated at two. See also Table 3 for grazing possibly fish as well) may be significant. Although
rates. replicates were taken and means and standard errors
were produced, no statistical tests are reported.
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location, study
type
Bittium Zostera Examines how the interaction between seagrass and epiphyte Neckles et Seasonal experiments were carried out under a
varium (G) marina. production, grazing and nutrient addition are affected by the time of al. 1993 randomized block pattern in mesocosms to examine the
Mitrella Chesapeake year. During spring and summer, epiphyte biomass increased with combined interactions of grazers, seagrass production
lunata (G) Bay, Virginia both removal of grazer and nutrient enrichment. In early summer, and epiphyte growth under ambient and 3×ambient
Idotea USA. with low ambient grazer densities, seagrass production was reduced nutrient regimes. Grazer densities approximated
baltica (I) Mesocosm only with grazers absent and under nutrient enrichment. During the ambient field conditions. This meant that grazer species
Erichsonella experiments autumn there was little epiphyte response and no seagrass response. and density changed significantly with season e.g. B.
attenuata (I) A conclusion is that the influence of epiphytes on seagrass production varium was present at densities of 4000 m−2 in early
Gammarus may be related to changing water temperatures and to seasonal
sp. (A) changes in macrophyte requirements for light and inorganic carbon. summer and 0 m−2 in autumn and spring. Data were
Ampithoe sp. analyzed using repeated measures analyses of variance
(A) with appropriate tests of statistical assumptions of the
analysis. In summary, a well designed experiment with
sound statistical analyses and interpretation.
Tripneustes Thalassia A study examining the impact of sea urchins on tropical seagrass and Klumpp et The study was carried out between OctoberDecember
gratilla (U) hemprichii. macroalgae, sea urchin dietary preferences, assimilation efficiencies. al. 1993 1991. Field surveys of seagrass and macroalgal species,
Salmacis Bolinao, Seagrass and sea urchin density varied between the five study sites. abundance were carried out using 20 0.5 × 0.5 m
sphaeroides Philippines. T. gratilla fed mostly on attached seagrass leaves while S. quadrats. Estimates of seagrass production derived
(U) Subtidal, sphaeroides was a generalist, consuming attached and detached from work to be published elsewhere. Sea urchin
sheltered seagrass leaves, red algae and algal coated sediment and rubble. surveys using 10 transects at each of 5 sites. Field
nearshore Some algae such as Sargassum were avoided. Absorption studies of sea urchin grazing using cages. Laboratory
sites. efficiencies of the sea urchins were above 70% for seagrass leaves studies of food preference used 4 aquaria with 5 sea
with epiphytes and were lower for epiphytefree seagrass. Grazing urchins in each, Absorption efficiency calculated using
rates were proportional to body size. The estimated annual grazing the material in the final 2 cm of intestine, There is no
impact was 24% of annual seagrass production but owing to large mention of using faeces actually excreted from the
changes in sea urchin population structure and density, the impact animals to calculate assimilation efficiencies,
varies from less than 5% to 100% depending on the time of year and
location. See also Table 3 for grazing rates.
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location, study
type
Idotea Zostera An investigation of the effects of grazing by the isopod, Williams & Quantitative study using both field and laboratory
resecata marina. Idotea, and inorganic nitrogen in the water column and in the Ruckelshaus experiments. Data analysis included analyses of variance and
(I) Intertidal sediment on the growth of seagrass and epiphytes. The 1993 although power calculations were included, testing of
meadow in results suggest that epiphytes have the potential to control statistical assumptions of the analyses, e.g. normality of error
Padilla Bay, seagrass growth when the dominant grazer, Idotea, is absent terms were not reported. Because epiphyte material consisted
Washington and when nitrogen in the water column is abundant and water mainly of diatoms biomass samples were not ashed and would
temperatures are suboptimal for seagrass growth. However, have contained small fauna (larger fauna were removed prior
this potential is not realised in the field because of the to weighing). Although experiments of grazing considered the
combined interaction of the above factors. The study effect of Idotea, other herbivores such as littorines and
highlights the importance of considering the combined effect amphipods were assumed to be in equal densities in all
of physical and biological factors when studying seagrass treatments (although this was not tested) and have similar
production. effects that were assumed to be negligible.
Idotea Zostera This study investigates the interaction of cyanobacteria, Neckles et al. Zostera grown in 20 microcosms Five replicates were used
baltica (I) marina. heterotrophic flagellates, nutrients and grazing on epiphytic 1994 for each partitioned into a 2 × 2 factorial design. treatment
Ampithoe Chesapeake diatoms, heterotrophic bacteria, isopods and amphipods. which were: Nutrient levels (ambient or enriched) and
sp. (A) Bay, Virginia Diatom abundance (cell counts) appeared to be unaffected by Grazing (present or absent). Data were analyzed by analyses
Gammarus USA. increased nutrients but reduced by grazing whereas addition of variance with appropriate tests of statistical assumptions,
sp. (A) Aquarium but were unaffected by grazers. cyanobacteria increased
experiments under nutrient Bacteria populations appear to be nutrient
limited and there was a strong correlation between
heterotrophic flagellates and bacteria. Heterotrophic
flagellates act as a link between heterotrophic bacteria and
higher trophic levels in seagrass epiphyte food webs. Ambient
grazer densities were used.
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location, study
type
Thalotia Posidonia A study using modified clear acrylic chambers to examine the relative Jernakoff & This quantitative field study used four
conica (G) sinuosa. impacts on periphyton and epiphytes by crustacean and gastropod grazers. Nielsen (in replicates of four chamber treatments;
Small Coastal Biomass of periphyton on seagrass was measured by ashfree dry weight press) crustaceans and gastropods, crustaceans
crustaceans meadows at 4 and chlorophyll. Impacts were measured 14, 21, 28 and 35 days after the alone, gastropods alone, no grazers, plus a
(A, I) m water depth, chambers were deployed. The results of the study are still very tentative: it control chamber treatment. Four sampling
Western appears that the molluscs had the greatest impact on the biomass of periods were measured. The majority of
Australia periphyton and epiphytes although it appears that crustacean grazing data were analyzed by analyses of variance
reduced the species composition. with appropriate tests of statistical
assumptions. Epiphyte species composition
data were analyzed by multivariate methods,
Pyrene Posidonia The study examined grazing and assimilation of seagrass, seagrass debris, Nielsen & Preliminary pilot studies and statistical power
bidentata (G) sinuosa. periphyton and large epiphytes by gastropod and amphipod grazers. Grazing Jernakoff analyses were used to adjust subsequent
Cantharidus Coastal waters rates were measured by biomass changes in grazed and ungrazed control (unpublished experiments on grazing. The data were
lepidus (G) at 4 m water treatments and assimilation efficiencies were measured by the biomass data) analyzed by analyses of variance with
Small depth. difference between food consumed and faeces excreted. Periphyton appropriate tests of statistical assumptions,
crustaceans Western (consisting of diatoms and algal propagules) were most easily consumed and
(A) Australia assimilated. The grazing rate and assimilation of periphyton by the
Aquaria amphipods was twice that of the gastropods. Grazing rates in summer were
experiments higher than in winter. The larger epiphytes Ulva, Cladophora and
Hypnea, were consumed by amphipods but not by gastropods. Seagrass
and seagrass debris were not consumed. See also Table 3 for grazing rates.
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Spatial and temporal patterns of abundance of seagrass epiphytes and their grazers
In this section we review the factors influencing the spatial and temporal patterns of abundance of seagrass epiphytes and their grazers. These factors operate over a
range of scales. For example, spatial differences may be apparent at the scale of a seagrass leaf, or along an environmental gradient. Temporal differences may be
important on diel or yearly scales. We examine these factors separately for the epiphytes and grazers and then in our final section we explore how their interaction
results in the overall pattern of epiphytes and grazers in seagrass meadows, and the implications of measuring them at different spatial and temporal scales.
Epiphytes
Although the diversity of seagrass epiphytes has been well documented (Humm 1964, Ducker et al. 1977, Harlin 1980, Cinelli et al. 1984, Heijs 1985a, Kendrick et
al. 1988, Borowitzka et al. 1990), there is relatively little information on their spatial and temporal patterns of abundance (Borowitzka & Lethbridge 1989). Spatial and
temporal patterns reflect the characteristics of the epiphytes as well as the morphology and life span of their seagrass host and the species pool within an area (Sullivan
1979, Heijs 1985a, Kendrick et al. 1988, Borowitzka et al. 1990). In addition, abiotic factors such as insolation (Jacobs et al. 1983), light (Cambridge et al. 1986),
nutrients (including increases attributed to anthropogenic activity) (Penhale 1977, Borum 1985, SandJensen et al. 1985, Orth & Moore 1986), salinity (Kendrick et al.
1988) and temperature (Heijs 1985c) will influence the occurrence and distribution of the epiphytes.
Spatial patterns
Spatial patterns can be considered on a number of scales, from very small (e.g. along a leaf and within a shoot) to very large (e.g. along latitudinal gradients). Although
the objectives of a particular study will largely determine which scale is appropriate, these may not always coincide with the scales that are important to the biology of
the epiphytes.
The smallest spatial scale that we discuss in this review is at the “within leaf” scale. The distribution of epiphytes along a leaf may be due to: (a) the relative age of
different sections of a leaf, (b) the chronological sequence of development of the leaf and (c) the orientation of the leaf to other parts of the seagrass shoot (Borowitzka
& Lethbridge 1989).
Small spatial scales

(i) relative age of leaf Seagrass leaves continually provide new surfaces that epiphytes can colonize because the leaves grow from a basal meristem and older leaves
are shed sequentially. There is, therefore, a gradient in age (and addition of new substrata available for colonization) from the younger leaf base to the older apex and
on a shoot, from the newly emerged inner leaf to older outer leaves. The life span of a leaf varies between seagrass species, and growth rates also vary with season and
habitat (Ott 1980, McComb et al. 1981, see also Table 1).
The abundance and diversity of epiphytes correlates with leaf age on some seagrasses,
Page 133
e.g. Amphibolis antarctica (Bramwell & Woelkerling 1984, Walker & Woelkerling 1988); Enhalus acoroides (Brouns & Heijs 1986); Heterozostera tasmanica
(Bulthuis & Woelkerling 1983a); Posidonia australis (Horner 1987); P. oceanica (Novak 1984, Casola et al. 1987); Phyllospadix (Willcocks 1982); Thalassia
hemprichii (Heijs 1985b,c); T. testudinum (Kitting 1984) and Zoster a marina (Borum & WiumAndersen 1980, Jacobs et al. 1983).
The greater abundance and diversity of epiphytes on older seagrass leaves could be due to greater time available for colonization and growth (Borowitzka &
Lethbridge 1989) or to the better conditions (e.g. light and water flow) that promote maturation and sexual reproduction (Heijs 1985b, Mazzella & Alberte 1986,
Horner 1987, Borowitzka & Lethbridge 1989). However, a decrease has also been observed in the biomass on the oldest leaves of some seagrasses just prior to
senescence. This may be due to space limitation by encrusting coralline forms, sloughing, or both (SandJensen 1977, Willcocks 1982, Horner 1987, Walker &
Woelkerling 1988).
Different assemblages of epiphytes may be found on different parts of a seagrass shoot. For example, the stem of Amphibolis lives much longer than the straplike
leaves and supports a more diverse epiphyte community of fastgrowing seasonal species as well as longerlived species.
(ii) chronological sequence of development When the new seagrass leaf tissue emerges from the protective sheath, bacteria and diatoms rapidly colonize its
surface (Novak 1984). Pennate diatoms, one of the most prominent forms, gradually develop into a mat covering the whole blade. With time, debris and diatom
frustules accumulate and form an amorphous crust on which filamentous green and red algae develop (Zostera marina Sieburth & Thomas 1973, Jacobs & Noten
1980, and Heterozostera tasmanica Bulthuis & Woelkerling 1983a). Kitting (1984) also found that coccoid and then naviculoid diatoms initially colonized the new
leaf blades of Thalassia testudinum. Later these were sequentially replaced or complemented by filamentous red algae, mixtures of encrusting coralline algae (mainly
Heteroderma and Dermatolithon sp.), which formed a thick crust and dominated the epiphytic community of the oldest leaves. Coralline algae are also known to be
early colonizers of other types of seagrasses (Humm 1964, Bramwell & Woelkerling 1984, Walker & Woelkerling 1988, Borowitzka & Lethbridge 1989).
(iii) orientation of the leaf surface The position across a seagrass leaf may also influence the distribution and diversity of epiphytes. Epiphytes appear to settle
preferentially on leaf margins in Zostera marina (Jacobs et al. 1983), Posidonia oceanica (Casola et al. 1987), and Thalassia hemprichii (Heijs 1985c). Epiphytes
such as Audouinella and Chondria not only settle but show preferential growth at leaf margins on the seagrass Thalassia hemprichii (Heijs 1985b). Heijs suggests
that, within the Ceramiales, the Rhodomelaceae also show a stronger affinity towards the leaf edge, whereas the Ceramiaceae are equally distributed on leaf edges and
faces. Although settlement may start preferentially at the edges, Borowitzka & Lethbridge (1989) also reported that encrusting coralline algae and ceramialean red
algae, which initially settled on the edge of Posidonia spp. leaves in Western Australia, gradually spread over the leaf blade. Thus, the distribution of epiphytes across a
seagrass leaf may also be a function of both the age of, and position on, the leaf.
Epiphytes may also be more common on one leaf surface than the other. Novak (1984) suggested that a lower density of macroepiphytes and a higher density of
diatoms and bacteria on the inner faces of Posidonia oceanica leaves may be due to water movement, although
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Heijs (1985b) suggested that different light regimes on the different leaf faces could produce this pattern. It is also possible that water movement is affected by the
shape and orientation of the seagrass leaves. However, Heijs (1985b) found no differences in the species or abundance of epiphytes on different leaf surfaces of the
tropical species Thalassia hemprichii in Papua New Guinea, and suggested that differences may only be apparent in species with longerlived leaves. Similarly,
Bramwell & Woelkerling (1984) found no difference in cover, density or fertility of encrusting algae Pneophyllum fosliella on the adaxial or abaxial side of
Amphibolis antarctica leaves in southern Australia.
The morphology of different sides of seagrass leaves may also influence epiphyte coverage. Borowitzka & Lethbridge (1989), describing unpublished work,
reported that epiphytes were more diverse on the inner, concave leaf surfaces than on the outer, convex surface of the seagrass Posidonia sinuosa. P. australis
growing in the same area had similar numbers of species on both sides of its flat leaves. However, as no measures of the variability of epiphyte diversity within and
between species and leaves were provided, the result might be due to a sampling artefact. Trautman (1990) also found the same patterns of epiphyte cover on P.
australis and P. sinuosa.
Larger spatial scales

It is not possible to review all larger spatial scales here, so we have chosen to review the effect of environmental gradients on epiphyte abundance. Two environmental
gradients common in temperate seagrass meadows are salinity and depth.
Salinity gradients occur in estuaries and embayments. A decrease in epiphyte diversity on stems of Amphibolis antarctica in Shark Bay, Western Australia,
correlated with an increase in salinity (Kendrick et al. 1988). The epiphytes could be grouped into three types: species that have high relative abundances in areas of
salinity greater than 50‰ but which occur throughout the salinity range; species that only occur in areas with salinities less than 50‰; and species that show no
relationship between relative abundances and salinity. Sixtysix species were recorded with members of the Rhodophyta (68%) dominating the flora, followed by the
Phaeophyta (23%) and the Chlorophyta (9%). On a percentage basis the community was similar to that reported for the southern Australian marine flora (Womersley
1981). A study of four species of encrusting coralline algae along the same salinity gradient in Shark Bay found no change in the species composition, although
abundance and reproductive development decreased with increasing salinity (Harlin et al. 1985).
Epiphyte diversity and biomass have been correlated with changes in water depth, which also correlated with changes in light, temperature and hydrodynamics
(Mazzella et al. 1989). The abundance and fertility of crustose epiphytes increased on deeper sublittoral stands of Heterozostera tasmanica and Amphibolis
antarctica in southeastern Australia (Bulthuis & Woelkerling 1983a, Bramwell & Woelkerling 1984) and Posidonia oceanica in the Mediterranean (Cinelli et al.
1984). In contrast, noncoralline epiphytes were negatively correlated with depth (Cinelli et al. 1984). These patterns may vary with season (Cinelli et al. 1984, Mazzella
et al. 1989).
Temporal patterns
Temporal patterns may be recognizable at a number of scales; in units of hourly, daily, seasonal and interannual changes. The appropriate scale will depend on the
biology of the epiphyte (e.g. growth, reproduction and settlement) and the purpose of the study. Changes in the microalgal community can be rapid, occurring over
hours or days, whereas change may
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be slower in the larger macroalgae (SandJensen & Borum 1991). Daily and hourly changes in microalgae may be detectable only by very finescale sampling, so
although they may be important in terms of periphyton dynamics, we will focus our discussion on seasonal patterns.
Seasonality affects abundance and diversity patterns of most epiphytes, but some do not show a seasonal pattern; they are present throughout the year (Brauner
1975) or occur so infrequently that patterns cannot be detected (Borowitzka & Lethbridge 1989). For example, Mazzella et al. (1989) found that about a third of the
epiphytes on Posidonia oceanica in the Mediterranean were present in both seasons. Most were rhodophytes, followed by phaeophytes and then a few chlorophytes.
Ducker et al. (1977) found a similar pattern on the seagrasses Amphibolis antarctica and A. griffithii in Australia.
Most epiphyte species peak in biomass in the warmer months but the timing of the peak varies between species (May et al. 1978). There are communities that peak
in spring (Humm 1964, Penhale 1977, Mazzella et al. 1989, Edgar 1990), communities that peak in summer (Wood 1959, May et al. 1978, Sullivan 1979, Jacobs &
Noten 1980, Bulthuis and Woelkerling 1983b, Heijs 1984, 1985a, Morgan & Kitting 1984, Brouns & Heijs 1986, Bell & Westoby 1987) and those that peak in late
summer and early autumn (Brauner 1975, Heijs 1984, 1985a, Silberstein et al. 1986, Borowitzka et al. 1990). Seasonal peaks may also depend on local
environmental conditions. For example, Edgar (1990) found a 2month difference between the peak abundances of coralline epiphytes on A. griffithii at two sites
35km apart and suggested that differences in water temperature might be responsible.
Seasonal patterns of some epiphyte species have also been closely linked with the seasonality of their host seagrasses. Penhale (1977) suggested that the growth of
the eelgrass Zostera marina influenced the growth of its epiphytes. Similarly, Jacobs & Noten (1980) suggest that changes in the diatom community are controlled by
the productivity of their eelgrass host through a direct nutrient link. Although a correlation was established in both of these studies, a causeandeffect relationship has
yet to be demonstrated. In particular, the likely possibility that both seagrasses and their epiphytes are controlled by the same external seasonal factors has yet to be
disproved.
In summary, the factors affecting epiphyte distribution operate at very small and very large spatial scales. Although some of the smaller scales may not be important
when assessing changes in epiphyte abundance or species composition over larger areas, they are relevant when designing sampling and survey methods. Given the
variability of epiphytes on the leaves, based on their position on the leaves and the age of the leaves, it is important to ensure that comparisons between studies fully
account for the effect of different sampling designs (sample equipment and technique, space and timescales of sampling). Standardized survey methods to address
spatial heterogeneity are particularly important along environmental gradients. Similarly, given the different seasonal patterns among species, care is also needed in
comparing studies of different epiphyte species sampled at different times of the year.
Grazers
Seagrass faunal communities can be divided into epifauna, infauna, epibenthic and sessile, according to their distribution within the meadow (Hutchings 1982). Motile
epifauna generally consist of amphipods, gastropods, isopods and freeliving polychaetes. Sessile epifauna are mainly hydrozoans, bryozoans and tubiculous
polychaetes; infauna are mainly burrowing
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amphipods and a variety of bivalves, polychaetes and decapod crustaceans; epibenthic species are mainly resident and transient fishes (Hutchings 1982, Howard et al.
1989, Hutchings et al. 1991). These divisions are convenient for researchers (Howard et al. 1989), but in practice the distribution of fauna within these classifications
may vary spatially and temporally (Greening & Livingston 1982, Howard 1987).
Amphipods and gastropods form a considerable subset of the epifauna and they are important grazers of seagrass epiphytes (Nagle 1968, Marsh 1973, Zimmerman
et al. 1979, Nicotri 1980, Howard 1982, van Montfrans et al. 1984, Virnstein et al. 1984, Virnstein & Howard 1987a). However, most studies of spatial and
temporal patterns of seagrass epifauna do not distinguish between grazers and other members of the community. This section, therefore, also reviews patterns of
epifauna while focusing on the grazer component of the epifauna.
Spatial patterns
As for epiphytes, spatial patterns of grazers can be considered on such small scales as along a seagrass leaf to the large scale environmental and latitudinal gradients.
Small spatial scales

Smallscale patterns in the distribution of epifauna occur among microhabitats within individual seagrass plants, e.g. the position along the leaf on zosteroid plants and
between leaves and stems on amphiboloid seagrasses. Nagle (1968) recognized three distributional patterns of epifauna on individual Zostera marina leaves: those that
increase in abundance up the leaf; those that decrease in abundance up the leaf; and those that exhibit no clear distributional pattern. For example, crustaceans such as
amphipods, ostracods and copepods were generally distributed with increasing abundance towards the tips of the leaves, although the amphipod Corophium
ascherusicum was more abundant at the base of the plants. Similarly, the snails Bittium and Caecum were located peripherally, whereas Crepidula and Littorina
were distributed basally, as were most polychaetes and nematodes. Amphipods such as Microdeutopus damnoniensisi, M. grylloptalpa and Cymadusa compta
were distributed near the base of epiphytes on Zostera marina, whereas Corophium acutum and C. insidiosum were located distally on epiphytes. These
distributions were maintained by aggressive interspecific interactions (Nagle 1968). Edgar & Robertson (1992) also identified amphipods that were more abundant
peripherally on epiphytes. Although these distributional patterns were observed during the day, they may have little relevance to grazing patterns because of nocturnal
migrations (see p. 138).
At a slightly larger scale, seagrassassociated epifauna show little hostspecificity, and “typical” communities associated with particular seagrass species are not
apparent. In eastern North American seagrass meadows, similar epifaunal suites have been collected from Halodule wrightii, Thalassia testudinum and
Syringodium filiforme meadows, despite the plants having different morphologies (Stoner 1983, Lewis 1987, Virnstein & Howard 1987a). Virnstein & Howard
(1987a) report an 82% similarity in species composition between meadows of these three seagrass species and a 75% similarity between these meadows and those of
the red alga Gracilaria sp. (Virnstein & Howard 1987b). The species composition of epifauna in adjacent Posidonia australis and Zostera capricorni meadows in
eastern Australia overlaps broadly (Young 1981), as it does in Amphibolis and Posidonia in Shark Bay in Western Australia (Wells et al. 1985).
Despite the faunal composition of seagrass species overlapping, there are generally quan
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titative differences in the faunal species’ distribution. For example, gastropod and amphipod species in southern Australian meadows of Amphibolis antarctica,
Posidonia angustifolia and P. sinuosa overlapped broadly but, based on faunal abundances, the Amphibolis and Posidonia suites were distinct: the gastropods
Cantharidus irisodontes and C. bellulus preferred the Amphibolis habitat, while the amphipods Cerapus abditus, Tryphosella orana, Parawaldeckia sp.,
Ericthonius sp. and Tethygeneia sp. and the gastropod Thalotia conica preferred the Posidonia habitat (Sergeev et al. 1988). Similarly, among southwestern
Australian epifaunal gastropods, the trochids Prothalotia lehmanni and Cantharidus pulcherrimus preferred Amphibolis griffithii meadows, while Thalotia conica
and Cantharidus lepidus preferred adjacent Posidonia australis meadows. The columbellid Pyrene bidentata was found among both seagrasses but occurred more
patchily within the Posidonia australis meadows (Nielsen 1987).
Many studies have shown that the abundance and diversity of epifauna tend to increase with the abundance of benthic macrophytes (Nagle 1968, Marsh 1973,
Nelson 1980a, Gore et al. 1981, Lewis & Hollingworth 1982, Virnstein et al. 1984, Lewis 1987). Earlier studies of spatial patterns of epifaunal distribution focused on
the role of seagrass biomass and surface area in structuring these assemblages (Stoner 1980a,b, Stoner & Lewis 1985). However, attempts to demonstrate linear
relationships between habitat variables such as macrophyte biomass and surface area and faunal community parameters have failed (Stoner 1980a,b, Virnstein et al.
1984, Stoner & Lewis 1985).
The relationship between faunal abundance and seagrass biomass only holds within a seagrass species and when comparing the faunal communities of different
seagrass species, plant morphology has a greater influence (Stoner 1980b). Consequently, comparisons of the fauna of communities of different seagrass species are
difficult to make and depend upon the particular spatial scale being considered. Stoner (1983) found that Thalassia meadows supported a greater abundance of
amphipods than did adjacent Halodule and Syringodium meadows when compared in terms of unit area of seabed. When the data were standardized against seagrass
biomass, there was no significant difference between the amphipod abundances in Thalassia and Halodule, but Syringodium supported a significantly greater
abundance. Similarly, Virnstein & Howard (1987a) found that gastropods were most abundant on Halodule and crustaceans on Thalassia, in terms of seabed area,
but when expressed as a function of seagrass biomass or surface area, abundance was greatest on Syringodium.
Even comparisons within a seagrass species indicate significant heterogeneity between meadows in the abundance of fauna (Collet et al. 1984). Collett et al. (1984)
found that of the species collected from numerous sites within a Posidonia australis meadow, 46% were collected from only single sites, and that the species
composition appeared to change dramatically over relatively small spatial scales.
Larger spatial scales

The abundance and composition of epifauna change along environmental gradients (e.g. depth, salinity), particularly in estuaries, and it is for these reasons that Virnstein
et al. (1984) suggest that local factors have an overriding influence on faunal distribution. Gambi et al. (1992) found a change in grazer guilds with depth in Posidonia
oceanica meadows: herbivores dominated the most shallow zones (1–3m), herbivores and detritivores dominated middepth zones (10m), and deposit feeders and
carnivores dominated deeper zones (15–25m). In summer, herbivorous gastropods and amphipods were more evenly distributed along a depth transect, which Gambi
et al. (1992) suggest is due to the more even distribution of the epiphyte biomass and to depthrelated growth rhythms of both the epiphytes and
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the seagrass. Similar effects of depth on amphipod guilds have been reported in Tasmanian benthic algal habitats, where variations in abundance were related to the
distribution of the benthic algae (Edgar 1983a).
The composition of grazer communities has also been linked with hydrodynamic gradients. Nagle (1968) found that the gastropod Bittium, a tallspired snail, was
found in areas where water action was gentle, whereas the more globose Lacuna was found in areas where the waves were stronger. In Australian Amphibolis
meadows, Edgar & Robertson (1992) found that freeliving amphipod genera such as Tethygeneia, Hyale, Gitanopsis and Cyproidea were easily swept off seagrass
leaf surfaces, whereas tubedwelling genera such as Cymadusa and Ampithoe were less susceptible to water movement.
Combinations of environmental gradients may also influence the spatial pattern of epifauna. Dugan & Livingston (1982) observed that a decline in species richness of
epifauna in Thalassia meadows coincided with times of environmental stress, such as low water temperature and salinity. Similarly Nelson et al. (1982) sampled
Halodule wrightii in Indian River Lagoon, Florida, at sites that represented an environmental gradient in terms of water temperature, salinity and water depth. From
their survey results, Nelson et al. (1982) concluded that variations in the species richness and abundance of amphipods were more likely related to levels of predation
than to environmental factors.
Latitudinal patterns
Latitudinal patterns in some taxa of seagrass epifauna have been reviewed by Virnstein et al. (1984). They found that relationships between latitude and the diversity
and abundance of epifauna were inconsistent: for example, the diversity of decapods and amphipods increased significantly with decreasing latitude, whereas isopod
and fish diversity show no latitudinal trends. However, they concluded that local factors such as habitat complexity and predation are of greater importance in
structuring epifaunal communities. Clearly, though, there are ecological parallels in the composition of faunal communities in different seagrass habitats over a range of
geographical regions (Nagle 1968, Marsh 1973, Nelson 1980a, Lewis & Hollingworth 1982, Virnstein et al. 1984).
Temporal patterns
Diel, seasonal and annual timescales are important and can be recognized in the distribution and abundance of epifaunal grazers.
Diel patterns
Vertical and horizontal migrations of seagrass fauna are usually a local, nocturnal phenomenon associated with feeding (e.g. Howard et al. 1989), avoidance of
epibenthic predators (e.g. Williams & Bynum 1972, Tranter et al. 1981, Fancett & Kimmerer 1985), reproduction (e.g. Williams & Bynum 1972, Robertson &
Howard 1978) or dispersal (e.g. Robertson & Howard 1978). However, some species remain in the seagrass canopy during the day and night, e.g. Prothalotia
lehmanni, which grazes over Amphibolis (Nielsen & Lethbridge 1989).
Several studies of seagrass faunal communities suggest that surveys based solely on diurnal collections may give an erroneous picture of the community (Greening &
Livingston 1982, Ward & Young 1982, Howard 1987). Many benthic fauna migrate vertically at night, either from the seagrass canopy to the water column, or from
the rhizome mat to the canopy or water column. However, no examples of downward vertical migration at night have been
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reported (Howard et al. 1989). Amphipods, cumaceans, polychaetes and copepods form part of the demersal zooplankton at night (Robertson & Howard 1978,
Alldredge & King 1980, Ohlhorst 1982). Gastropod molluscs migrate vertically along Posidonia leaf surfaces at night (Russo et al. 1984, Nielsen & Lethbridge
1989). Such migrations affect the spatial distribution of seagrass fauna over relatively short timescales, so must be considered when designing sampling programmes.
A similar total number of fauna species and individuals were collected during the day and at night in mixed Thalassia, Syringodium and Halodule meadows
(Greening & Livingston 1982). However, the abundances of many species were significantly different; for example, decapod crustaceans, which emerge from the
sediment at night, were more abundant in nocturnal samples. The patterns were not consistent among the fauna, however, so generalizations on the diel patterns of
different taxa cannot be made from this study. In Halodule wrightii in the Indian River lagoon, Florida, Heck (1977) also found higher nocturnal abundances of a
variety of decapod crustaceans; during the day species such as penaeid prawns and portunid crabs burrowed into the sediments and caridean shrimps remained near
the sediment surface.
The abundance of seagrass fauna was greater by night than by day in Howard’s (1987) study in H. wrightii meadows in the Indian River Lagoon, Florida, with the
crustaceans averaging a twofold increase in nocturnal samples. Of the ten most abundant species collected, seven were significantly more abundant at night, including
the gastropods Crepidula convexa, Bittium varium, Cerithium muscarum, Modulus modulus and the amphipod Cymadusa compta. In Puerto Rican Thalassia
meadows, Bauer (1985) also found that while the species composition was similar, abundances were considerably higher at night than in the day. Diel differences in
abundance were greatest for species that stay in the sediments by day (e.g. Processa spp.) and lowest for inhabitants of the canopy (e.g. Hippolyte and Latreutes).
At night, gastropods migrate vertically upwards along seagrass leaves, as did columbellid and phasianellid gastropods in a Posidonia oceanica meadow studied by
Russo et al. (1984). In southwestern Australian meadows the trochid Thalotia conica and the columbellid Pyrene bidentata migrate up the leaves of Posidonia
australis at night, but no such migrations were observed in nearby Amphibolis griffithii meadows (Nielsen & Lethbridge 1989). Possibly the morphology of
Amphibolis, which has leaf clusters at the end of perennial lignified stems, does not facilitate such migrations. However, Edgar & Robertson (1992) report that the
gastropods Microdiscula charopa, Tricolia sp., Eatoniella australiensis and Sinezona sp. were dependent on the presence of both seagrass leaf and epiphyte
habitats. They reside among epiphytes by day and migrate onto the seagrass leaf by night, possibly to feed. Kitting (1984) found that the grazing activities of Anachis
avara and Bittium varium were more frequent at night, when the snails were in the seagrass canopy, than by day, when they were near the sediments.
Longterm changes in the distribution of epifauna among microhabitats have received little attention, although Robertson & Mann (1982) report a seasonal change in
the distribution of Littorina neglecta on Zostera marina leaves. During the winter Littorina neglecta is found closer to the sediment surface, possibly to avoid winter
ice (Table 2).
Seasonal patterns
Epifaunal species may undergo large cyclical fluctuations in abundance (Marsh 1973, Nelson 1980b, Stoner 1980c, Dugan & Livingston 1982, Edgar 1990). The
abundance of epifauna usually peaks in springsummer, with a corresponding decrease in winter (Marsh 1976,
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Heck 1977, Stoner 1980c, Nielsen 1987). Seasonal fluctuations in grazer abundance can range over orders of magnitude, e.g. densities of amphipods can go from
about 10000m−2 to over 100000m−2 in Amphibolis meadows (Edgar 1990).
Although spatial patterns of epifaunal abundance and distribution are strongly influenced by the complexity of the habitat (seagrass biomass, architecture, attached
epiphyte load), it is unclear whether seasonal patterns are similarly influenced. Nagle (1968) and Marsh (1973) report that in the Northern Hemisphere, Zostera
declines during winter and that epifaunal abundance generally follows this pattern. Nagle (1968) recorded summer densities of 9000 amphipods m−2 that when
extrapolated to seabed area, increased during winter to 37500m−2, albeit on isolated clumps of Zostera. In Florida, Dugan & Livingston (1982) found that macro
invertebrate abundance became positively correlated with increased macrophyte production during spring and summer and that annual patterns of variation were
detectable for at least as long as 7 years.
Conversely, many studies point out the importance of external forces in seasonal patterns of epifaunal abundance. Stoner (1980c) suggests that, although macrophyte
biomass governed the spatial distribution of amphipods, seasonality of benthic macrophytes did not appear to be important in explaining temporal variability in
amphipod abundance, which was related more to the abundance of predators. Similarly, Nelson (1979) and Nelson et al. (1982) report that amphipods in Halodule
wrightii meadows were seasonally more abundant between November and May (late autumn to late spring), which inversely related more closely to seasonal
abundances of fishes and decapod predators than to seasonal patterns of seagrass biomass. These patterns were consistent between different years and between
different sites within any given year. Robertson & Mann (1982) report that 2+yearold individuals of Littorina neglecta experience a high mortality due to fish
predators.
Seasonality of reproduction and recruitment may also influence the temporal abundance of seagrass epifauna. In eastern Australian Zostera capricorni and
Posidonia australis meadows, annual patterns of decapod fauna appeared to be related to seasonal patterns of recruitment and mortality (Young 1981). Similarly,
Robertson & Mann (1982) and Edgar (1990) found that populations of epifaunal gastropods peak in springsummer during periods of high recruitment, but decline
quickly due to high juvenile mortality. Annual patterns of abundance of Western Australian epifaunal gastropods may be related to the breeding periodicity and mode of
reproduction of these species (Nielsen 1987). For example, Thalotia conica and Prothalotia lehmanni, which are broadcast spawners that reproduce through spring
and summer, show seasonal cycles of abundance, whereas Pyrene bidentata, which lays egg capsules, with about 20 embryos in each, throughout the year has a more
stable yearround abundance. Stoner (1980c) working in Florida, found that seasonal patterns of amphipod abundance were related to their reproductive pattern.
However, Nelson (1979) working in north Carolina, concluded that physical factors and reproductive seasonality have little to do with overall seasonal variations in
amphipod abundance, as most of the species studied reproduce throughout the year.
Few studies have looked at the influence of the seasonal availability of food on abundance and distribution of epifauna. Heck & Wetstone (1977) suggest food
resources are probably not a limiting factor for mobile epifauna. However, Robertson & Mann (1982) found that Littorina neglecta abundance in Zostera marina
peaked in midsummer at 45000m−2, but declined rapidly because food resource limitations caused a high mortality of recruits. Littorina neglecta controlled the
growth of epiphytes during this time. Edgar (1990) also reports that juvenile mortality of Cantharidus lepidus in Amphibolis meadows may be due to food resource
limitations. In contrast, Nelson (1979) found no evidence of food limitations
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or competition for food among amphipods in Zostera meadows, as peaks in the abundance of amphipods coincided, despite a large overlap of diets. Additionally,
Schneider & Mann (1991) suggest that there is a surplus of periphyton and filamentous algae on Canadian Z. marina meadows. Food quality may be more important
than food quantity because Jackson (1972) observed that the diversity of bivalves in Jamaican Thalassia meadows was not correlated with the total quantity of food
available, but was correlated with the variety of available food.
This section has emphasized the relevance of scale to the study of the biology of grazers at the individual, the population and the community level. Although large
scale patterns such as correlations along environmental gradients do exist, local influences such as hydrodynamics, predation and habitat complexity probably have a
stronger influence on the distribution and abundance of grazers in seagrass meadows. Therefore, sampling of these populations requires particular recognition of
heterogeneity in space and time. This is particularly important when comparing different studies and extrapolating from results.
Grazerepiphyte interactions
The previous sections of this review highlight the multitude of factors that could influence the magnitude, direction and outcome of the interactions of grazers and
seagrass epiphytes. The biological factors include the epiphytes’ physical and chemical defences to prevent grazing (e.g. Steneck & Watling 1982, Hay et al. 1987,
Mazzella et al. 1992), feeding mechanisms of grazers (e.g. Howard 1982, Nielsen & Lethbridge 1989), sizeselective and speciesselective grazing preferences (e.g.
van Montfrans et al. 1982, Peduzzi 1987), resistance to digestion (e.g. Jernakoff 1985, Santelices & Ugarte 1987), growth and grazing rates of epiphytes (e.g.
Hootsmans & Vermaat 1985, Mazzella et al. 1992), habitat preferences (e.g. Edgar & Robertson 1992), predation on grazers (e.g. Nelson 1979, Virnstein 1979,
Heck & Orth 1980, Stoner 1980c, Orth et al. 1984) and the vagaries of colonization, settlement and seasonal patterns (e.g. Edgar 1992, Neckles et al. 1993). The
physical factors include temperature and water movement (e.g. Borowitzka & Lethbridge 1989), water depth (Cinelli et al. 1984), habitat complexity (Nelson 1979)
and nutrient enrichment (Neckles et al. 1993, 1994, Williams & Ruckelshaus 1993).
Grazerepiphyte interactions can be partitioned into the effect of grazing on the presence and abundance of epiphytes, and the effect of epiphytes as a food resource
and habitat on the presence and abundance of grazers.
Effect of grazers on epiphytes
Removal of epiphyte biomass

Grazers can reduce epiphyte biomass on seagrass leaves (Zimmerman et al. 1979, Howard 1982, Robertson & Mann 1982, van Montfrans et al. 1982, Orth & van
Montfrans 1984, van Montfrans et al. 1984, Hootsmans & Vermaat 1985, Howard & Short 1986, Borum 1987, Peduzzi 1987, Nielsen & Lethbridge 1989, Mazzella
et al. 1992, Neckles et al. 1993, Williams & Ruckleshaus 1993; see Table 2). Their effects on the biomass depend on their species and density (Kitting 1984, Howard
& Short 1986, Hootsmans & Vermaat 1985, Peduzzi 1987, Mazzella & Russo 1989), and on their consumption rates and dietary prefer
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ences (Borum 1987, Mazzella & Russo 1989; Nielsen & Lethbridge 1989), as well as the species and growth rates of the epiphytes (e.g. Borum 1987, Peduzzi 1987).
Small grazers such as amphipods have a variety of feeding strategies and food preferences (Borum 1987, Duffy 1990); some even feed on the macroalgal host as
well as epiphytes (Duffy 1990). Consumption of a seagrass host, however, is less likely, as seagrasses are not generally a preferred food item (Nicotri 1980). The roles
of amphipods may often be understated because numbers are generally kept below the carrying capacity by predation (Young & Young 1978, Nelson 1979, Stoner
1980c, Duffy 1990). However, the effect of amphipods when predation is low has been inferred by Tegner & Dayton (1987) working in southern Californian kelp
beds. There, low fish abundance, as a result of El Niño events, correlated with an increase in amphipod numbers and the destruction of kelp communities caused by
their excessive grazing. There have been no studies to date that report a similar impact on seagrass meadows.
Most studies of the effect of grazing on the epiphyte biomass have concentrated on species of Zostera, Thalassia, Halodule, Posidonia or Amphibolis (Tables 1
and 2, see pp. 112, 114). Furthermore, most of these studies concentrated on grazer diets or the removal of periphyton rather than filamentous epiphytes (e.g. Borum
1987, however see Zimmerman et al. 1979, Robertson & Mann 1982).
Studies of grazers in Australian temperate seagrasses include those of Howard (1982) and Nielsen & Lethbridge (1989). Howard (1982) studied grazing of diatoms
(Cocconeis scutellum, Synedra spp., Mastogloia sp. and Nitzschia sp.) by amphipods (Tethygeneia nalgo and Paradexamine churinga) on Heterozostera
tasmanica in southeastern Australia. His descriptive study compared epiphyte abundance and amphipod feeding in an aquarium with an ungrazed control. Howard
(1982) concluded that, because of its feeding morphology, Tethygeneia nalgo probably plays a major role in removing fouling material from the seagrass, but that
neither species of amphipod ate live seagrass material. From an analysis of the gut contents of prosobranch gastropods (Prothalotia lehmanni, Pyrene bidentata and
Thalotia conica) in Amphibolis griffithii in Western Australia, Nielsen & Lethbridge (1989) inferred that living epiphytes are a more important food source than
detritus, but did not comment on the effect of gastropods on epiphyte growth.
Apart from removing algal biomass, grazers can also remove detritus and other particulate matter falling on seagrass leaves. In a study of grazing by the gastropod
Littorina littorea in eelgrass meadows of Nova Scotia, Robertson & Mann (1982) found that, not only did the snails remove filamentous algae and diatoms, but they
also reduced the amount of detritus accumulating on the leaves. Their activities had positive effects on the seagrass’s growth.
Grazing and epiphyte production

The grazing rates and epiphyte growth of some seagrassassociated species have been estimated (see Table 3). Van Montfrans et al. (1984) concluded that
consumption of micro and macroalgal seagrass epiphytes (which are responsible for between 20 and 60% of production of the meadows, Orth (1992)) is probably
substantial, because while most epiphytes colonize and turnover rapidly, their actual abundance is far below the maximum potential. Harlin (1980) also suggested that
the normally low level of the epiphyte biomass may be due to the presence of grazers, and that the overall productivity of the epiphytes is probably high because their
turnover times are short. This view is shared by Peduzzi (1987), who suggested that trochid gastropods grazing on Posidonia oceanica in Italy removed 40% of the
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Table 3 Rates of grazing (units quoted as in original reference; n/a grazer density and size not available or not applicable when individuals were used in
feeding trials), m= male; f=female.
Group Species Grazer density/size Macrophyte Grazing rate Units Reference

Amphipods Cymadusa compta Gammarus n/a Epiphytes on Thalassia testudinum, 0.87 mg ingested mg Zimmerman et al.
mucronatus Melita nitida Halodule wrightii and Syringodium 0.90 amphipod−1 day−1. (mean 1979
Grandidierella bonnieroides filiforme 1.05 values for epiphytes on all
0.35 three species)
Ampithoe mea n/a Sargassum muticum 4.4±0.63 Mean±SE mg (ww) Norton &
amphipod−1 day−1 Benson 1983
Allorchestes compressa n/a Ecklonia radiata (holdfast tissue) 0.011 g g−1 day−1. Robertson &
Ecklonia radiata (eroding tips) 1.811 Lucas 1983
Ampithoe longimani n/a Dictyota dichotoma and three 19 mg eaten per 19 Hay et al. 1987
epiphytic species: Cladophora 36 amphipods in 46 hours.
ruchingeri, Polysiphonia harveyi, 24
Enteromorpha linziformis 18
Ampithoe marcuzii n/a Sargassum 2.5 mg amphipod−1 day−1 Duffy 1990
Punctaria 0.5
Enteromorpha 0.25
Ectocarpus 0
Codium 0
Amphipod assemblage consisting of: 40 adult individuals Epiphytes of Posidonia sinuosa Mean+SE mg AFDW
Tethygeneia nalgo, per 10 cm × 10 Periphyton (summer) 1.727±0.287 food mg AFDW grazer
−1
Nielsen &
Paradexamine churinga, Hyale cm×40 cm aquaria Periphyton (winter) 0.740±0.161 day−1 Jernakoff
rubra and Ampithoe sp. Ulva (summer) 0.518±0.122 (unpublished
Ulva (winter) −0.073±0.152 data)
Cladophora (summer) 0.118±0.043
Hypnea (winter) 0.537±0.043
Isopods Idotea baltica n/a Ascophyllum nodosum and epiphytes 1 mg day−1 (for adults 20– Strong & Daborn
25 mm length) 1979
2559.4 Joules for males
2528.4 Joules for females (based
on 12.9835 J mg (DW) A.
nodosum−1)
Idotea baltica n/a 10 preferred species only mg ind.−1 day−1 Nicotri 1980
Codium 8.1
Porphyra 5.9
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Spongomorpha 3.8
Enteromorpha 3.3
Petalonia 3.3
Dicty siphon 3.1
Ulva 2.9
Sargassum 2.8
Scytosiphon 2.3
Hypnea 1.8
Idotea baltica n/a epiphyte Enteromorpha on Chondrus 0.04 g−1 h−1. (calculated from Shacklock & Doyle
crispus data given) 1983
Idotea chelipes ≈natural epiphytes on Z. marina 1.78 (±0.27) Mean+SD g g−1 day−1, Hootsmans &
AFDW Vermaat 1985
Idotea baltica natural density epiphyte species on Z. marina not In chlorophyll In units per day Borum 1987
I. viridis separated i.e. combined. units only
Sphareoma
rugicuda
Theoduxus
fluviatilis
densities n/a mean sizes Epiphytes on Fucus Mean ± SD mg Fucus 10 mg Salemaa 1987
are as follows: Idotea−1 day−1,
Idotea baltica (m) 10.9 mg 1.1±0.7
Idotea baltica (f) 7.9 mg 2.6±1.9
Idotea chelipes 3.0 mg 1.6±1.4
(m) 2.1 mg 3.5±2.2
Idotea chelipes 6.9 mg 1.4±0.9
(f) 4.7 mg 3.3+3.3
Idotea granulosa
(m)
Idotea granulosa
(f)
Decapods Palaemonetes n/a epiphytes on H. wrightii 0.185–0.190 mg epiphyte mg shrimp−1 Morgan 1980
pugio
h−1.
Molluscs Bittium varium n/a periphyton on Z. marina 62.7% percentage removal van Montfrans et al.
1982
Littorina littorea ≈ natural epiphytes on Z. marina 0.097 (0.048) g g−1 day−1, AFDW (SD) Hootsmans &
Hydrobia ulvae 0.859 (0.108) Vermaat 1985
Littorina saxatilis natural density epiphyte species on Z. marina not In chlorophyll In units per day Borum 1987
Hydrobia sp. separated, i.e. combined. units only
Jujubinus striatus n/a diatoms 4.08 (+2.86) Mean+SD Peduzzi 1987
green algal film 3.70 (+2.03) μg (organic carbon).
Gibbula diatoms 1.78 (+1.37) mg grazer (total ww)−1
umbilicaris green algal film 1.72 (+1.42)
day−1.
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seagrass debris 0.61 (±0.21)
macroepiphytes 0.59 (+0.34)
Strombus mutabilis 5.4–3.2m−2 periphyton on Enhalus acoroides and 1.3–18.1 mg (AFDW). Klumpp et al. 1992
Cerithium tenellum Cymodocea serrulata 5.1–9.8 grazer−1 m−2
0.9–8.3 m−2
Pyrene bidentata Ten individuals per epiphytes on Posidonia sinuosa Mean ± SE mg Nielsen &
10 cm × 10 cm×40 Periphyton (summer) 0.133±0.058 AFDW food mg Jernakoff
cm aquaria Periphyton (winter) 0.029±0.010 AFDW grazer
−1 (unpublished data).
Ulva sp. (summer) 0.027±0.045 day−1
Ulva sp. (winter) 0.018±0.007
Cladophora sp (summer) −0.002 ±
Hypnea sp (winter) 0.003
−0.004±0.003
Cantharidus lepidus Periphyton (summer) 0.450±0.142
Periphyton (winter) 0.107±0.014
Ulva sp. (summer) 0.035±0.150
Ulva sp. (winter) 0.041 ±0.011
Cladophora sp. (summer) −0.006±0.004
Hypnea sp. (winter −0.004±0.003
Echinoids Paracentrotus lividus 8.0 m−2 Posidonia oceanica 0.00499 g shoot−1 day−1 Kirkman & Young
1981
Lytechinus n/a mixed diet based on availability not food 5.28 cm3 material day−1. Vadas et al. 1982
variegatus preference, T. testudinum constituted 17% 0.66
of diet with the major component being 0.192 g (DW) day−1,
detritus and sediment organic DW day−1,
Lytechinus n/a Thalassia testudinum 9 kg m−2 Valentine & Heck
variegatus 1991
Tripneustes gratilla 0.9–4.2 m−2 Thalassia hemprichii and epiphytes 0.56W0.34 g wet weight per Klumpp et al. 1993
Salmacis (tropical) urchin
sphaeroides 0.17W0.53
Oligochaetes and Gleotrichia n/a epiphytes on freshwater algae 4.23 mg algae (DW) mg Cattaneo 1983
chironomids Coleochaete spp. grazer (DW)−1
(freshwater)
day−1,
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epiphyte primary production (based on the production rates of Mazzella & Ott (1984) and grazing rates recorded in the laboratory). Peduzzi’s (1987) study and the
complementary work of Peduzzi & Herndl (1986) and Mazzella & Russo (1989) are some of the few attempts to synthesize the effects of epiphyte grazing on the
seagrass community. These authors found that the absorption efficiency of Gibbula umbilicaris varied with the diet: snails eating macroepiphytes lost up to 80% of
the ingested carbon in the faecal material, whereas when eating microflora they lost only 13–23%.
Change in species composition

Grazers can change the structure of a community in two ways. First, selective feeding can remove some species from an area, leaving the less preferred species.
Secondly, nonselective grazing removes competitive dominants (which are usually the fastergrowing species), allowing the slowergrowing, less competitive species to
establish (Steneck 1983, Stenecketal. 1991).
Selective feeding has been demonstrated in studies on seagrass epiphytes. Removal by Bittium varium of periphyton and loosely adhering diatoms Nitzschia and
Amphora on Zostera marina led to the community being dominated by the more tightly adhering diatom Cocconeis scutellum (van Montfrans et al. 1982). In this
study, Bittium also selected senescent rather than photosynthesizing diatoms. Similarly, Jacobs et al. (1983) suggested that the dominance of crustose red algae on
Zostera marina in France was due to trochid gastropods having grazed the more easily removed algae. However, the authors did not investigate grazing experimentally
or sample the grazers. There were also qualitative differences between grazed and ungrazed treatments of Halodule wrightii in experiments conducted in mesocosms
by Howard & Short (1986). Ungrazed leaves were initially colonized by filamentous algae that were replaced by diatoms and particulate material as the experiment
progressed. In contrast, leaves with grazers remained free of particulate material but large epiphytes such as Ulva and Enteromorpha appeared near the tips and
encrusting coralline algae were conspicuous. In Australian temperate seagrass meadows, Nielsen & Lethbridge (1989) found that at some times of the year, some
epiphyte species were more abundant in the diet of gastropods than would be expected from the natural abundance of the epiphytes on leaves. They inferred that the
gastropods were feeding preferentially, which might influence the natural species composition of epiphytes on the seagrasses.
Changes in species composition may also be due to differential digestion of propagules. Some algae, particularly the opportunistic species, can survive the digestion
process and grazing may stimulate spore release (Jernakoff 1985, Buschmann & Santelices 1987, Santelices & Ugarte 1987). These authors suggest that this might be
an important mechanism for dispersing, redistributing and even increasing the number of macroalgal propagules being dispersed. We are not aware of any studies
investigating this phenomenon in seagrass meadows.
Filter feeders
Filter feeders process a large volume of water to remove particulate material from the water column. Although filterfeeding molluscs are usually considered to be
herbivores, filtering phytoplankton as food (Bayne & Hawkins 1992), they are not generally considered to be grazers of seagrass epiphytes. They do graze plankton,
but we know of no studies of whether they graze on the settling propagules of seagrass epiphytes. However, Doering & Oviatt (1986) thought that filter feeders may
show a preference for large phytoplankton that sink
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rapidly, such as diatoms, rather than smaller phytoplankton species; possibly propagules of seagrass epiphytes would also fall into that category. There is also evidence
to suggest that when propagules are numerically abundant, they are not only ingested but also digested by filter feeders (Santelices & Martinez 1988), and are possibly
seasonally significant in their diets.
If filter feeders are very abundant, they may have localized effects. In shallow coastal seagrass meadows the volume of water they filter is unknown, but filtration
rates of 21h−1 g−1 have been calculated by Poore & Rainer (1974). Although the rate of filtering might be high, the proportion of filtered material rejected might also be
as high as 80–90% (Tenore et al. 1973, Sornin et al. 1988).
Effect of epiphytes on grazers

Epiphytes can affect grazers in several ways: they can be a food source (Orth & van Montfrans 1984, Mazzella et al. 1992), a habitat (Heck & Orth 1980, Edgar
1992), or a shelter from predation (Virnstein 1979, Holmlund et al. 1990). These three factors will influence the distribution and abundance of grazers in seagrass
meadows (Orth 1992).
Food
Several studies suggest that food may be a limiting resource for grazers, although the link has not been fully demonstrated (Robertson & Mann 1982, Edgar &
Robertson 1992). In experimental chambers enclosing Zostera marina, in Nova Scotia, there were significantly fewer diatoms and epiphytes after 26 days where the
snail Littorina neglecta was grazing compared with ungrazed controls (Robertson & Mann 1982). This study was carried out in summer in very shallow water (0.75m
at low tide). Robertson & Mann (1982) monitored percentage cover rather than the biomass of epiphytes, on the assumption that changes in cover were indicative of
biomass changes. During winter, both the cover of epiphytes and the abundance of snails decreased. After the ice thaw, both increased rapidly. Robertson & Mann
(1982) suggested that the increase in snails was a response to the increased food supply. However, it could equally well have been due to, for example, the higher
water temperatures. Support for a food limitation hypothesis comes from a study by Edgar (1990) on infaunal animals in Western Australia. He concluded that the
seasonal fluctuations in abundance of epifauna in Amphibolis antarctica and A. griffithii were partly attributable to food limitation. The epifauna increased rapidly in
enclosures where five times the normal amount of food per animal was made available.
There appear to be no studies of grazers of seagrass epiphytes that demonstrate that a change in food supply results in a shift in the allocation of resources for
reproduction. A correlation between the seasonal abundance of macroalgae and a change in the size and number of eggs per brood of amphipods has, however, been
described by Skadsheim (1984) and Kusano & Kusano (1988).
Habitat
Most studies of marine macrophytes have examined the effect of habitat complexity on macrofauna and fishes (see Bell & Pollard 1989 for a review). In addition to
supporting conspicuous herbivores, seaweeds characteristically support small invertebrates, which may
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achieve high densities. The mechanisms by which epiphytic algae support large numbers of epifauna have not been established. However, the influence of seagrass
epiphytes on the abundance and diversity of motile seagrass epifauna has been demonstrated (Lewis & Hollingsworth 1982) and in particular for meiofaunal
harpacticoid copepods by Hall & Bell (1988) and for amphipods by Heck & Orth (1980), Stoner (1980a,b,c), Edgar (1992), Stoner & Lewis (1985), and Edgar &
Robertson (1992).
The attributes of epiphytes that are used to describe habitat complexity and its effects on fauna are; biomass (Stoner 1980b, Lewis 1984), surface area (Lewis &
Stoner 1983), length, width and complexity of branches (Edgar 1983b). However, Lewis (1987), in a study of crustacean epifauna on Thalassia testudinum and
macroalgae in the Gulf of Mexico, found no single morphological feature of plants that was an adequate predictor of the abundance or species richness of the fauna.
Although abundances varied, there was a similar suite of crustaceans on all macrophytes, despite differences in the shape, architecture and complexity of the plants.
Habitat choice and the effects of epiphytes on fauna may be complex, and vary with particular groups of fauna and particular species, the morphology of the alga
(microalgae, encrusting, filamentous or fleshy), the kind of resource (food, habitat or shelter) and on the ability of the epifauna to utilize the resource (Stoner 1980a, Hall
& Bell 1988).
There was broad overlap in the dominant amphipod, isopod, tanaid and decapod species among Halodule wrightii, Thalassia testudinum and Syringodium
filiforme meadows and the epiphytic red alga Laurencia poitei, despite differences in plant morphology (Lewis 1987). When expressed in terms of macrophyte
biomass, epifaunal abundance was highest on L. poitei. In the same mixedseagrass meadows, Virnstein & Howard (1987b) found that gastropods were more
abundant on seagrass leaves but peracarid crustaceans were more abundant on the red drift alga Gracilaria sp. In Canadian Zostera marina meadows the gastropod
Mitrella lunata and the amphipods Corophium insidiosum and Gammarus mucronatus prefer coarse, branching algal habitats, whereas the amphipod Ampithoe
sp. prefers the canopy (Schneider & Mann 1991). Virnstein & Howard (1987b) also suggest that drift algae may provide better shelter and food resources than the
host seagrass. The role of drift algae in providing shelter for a variety of taxa has been documented (Gore et al. 1981, Kulczycki et al. 1981), and Coen et al. (1981)
suggest that drift red algae may be a limited resource for which epifauna possibly compete.
Stoner (1985) observed that amphipods and tanaids were more abundant on stands of the alga Penicillus capitatus than in the surrounding sparse Halodule
meadow. He suggested that this alga offers an “island” refuge that provides motile epifauna with a superior habitat to the surrounding seagrass meadow. The macroalga
Pelvetia fastigata may provide a similar island refuge for epifauna (Gunnill 1982), and benthic algal habitats have been found to support dense assemblages of
epifaunal amphipods (Fenwick 1976, Edgar 1983a, Russo 1989). In Australia, Edgar & Robertson (1992) identified distinct microhabitat preferences among epifaunal
amphipods in a mixed Amphibolis antarctica/A. griffithii meadow by manipulating epiphyte and leaf densities within the canopy. Freeliving amphipods such as
Hyale, Tethygeneia, Gitanopsis, Paradexamine and Cyproidea were associated with epiphytes in the seagrass canopy and declined in abundance when epiphytes
were removed.
Epiphyte and faunal abundance on Thalassia testudinum in Barbados were positively correlated (Lewis & Hollingworth 1982). Similarly the density of harpacticoid
copepods and the biomass of the brown alga Giffordia mitchelliae, the dominant epiphyte, had a significant, positive association in Thalassia testudinum meadows
in Florida (Hall & Bell 1988). Similar rates of colonization of artificial and natural seagrass led these authors to suggest that fauna were selecting the physical attributes
of epiphytes, rather than responding to biotic interac
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tions. However, the abundance of fishes and large decapods in Zostera capricorni habitats in southeastern Australia were found generally to decrease during a bloom
of Giffordia mitchelliae (Bell & Westoby 1987). The epiphytes may have lowered the seagrass canopy and reduced the space under the canopy, hampering the
movements of the fishes.
Edgar & Robertson (1992) found that reduction in epiphyte abundance resulted in a corresponding reduction in epifaunal (including grazer) abundance. By excluding
predators, the authors demonstrated that predators did not cause the decline, but they were unable to determine whether the decline was related to the epifauna using
the epiphytes as habitat or as a food.
Shelter from predation

The role of seagrass epiphytes (and seagrasses) in providing shelter for small grazers from predation is well known (Nelson 1979, Stoner 1980a, Robertson 1984,
Stoner & Lewis 1985). Most research has concentrated on predation of crustacean grazers by fishes. Nelson (1979), Heck & Thoman (1981), Stoner (1980a,c
1983) Edgar (1983c) and Orth (1992) suggest that predation is reduced by macrophyte complexity and that the relationship between habitat complexity and predation
success is nonlinear. However, Holmlund et al. (1990) have reported that, contrary to popular dogma, predation on amphipods was higher on complex, highly
branched epiphytes such as species of Hypnea than on simpler sheetlike forms such as Ulva. They suggested that amphipods were more visible on branched algae
because of greater background contrast and that omnivorous predators preferred foraging among complex algae. Hacker & Steneck (1990) suggest that the body size
of amphipods may influence their choice of habitat and shelter from predation because small amphipods preferred highly branched algae, while larger amphipods
preferred the more simple, thalloidshaped algae.
Effect of grazerepiphyte interactions on seagrasses

Many studies and reviews have suggested that the grazing of epiphytes has a positive effect on the productivity of seagrass by reducing competition from the epiphytes
(e.g. van Montfrans et al. 1982, 1984, Shaddock & Doyle 1983, Orth 1992, Neckles et al. 1993, Williams & Ruckleshaus 1993; Table 2). The amount of light
available for photosynthesis is generally believed to be the major factor responsible for epiphytes outcompeting seagrasses (Penhale & Smith 1977, Mazzella &
Alberte 1986). The possibility that epiphytes outcompete seagrass for water column nutrients has also been suggested (Williams & Ruckelshaus 1993). However,
seagrasses are able to utilize nutrients from the sediment that would be unavailable to epiphytes (Williams & Ruckelshaus 1993). These authors found that although this
alternative source of nutrients was available to Zostera marina, the seagrass was still nutrientlimited at ambient nutrient levels. They concluded from laboratory
experiments that the isopod Idotea resecata controlled epiphyte growth and in its absence epiphytes controlled Zostera production.
Other laboratory studies have experimentally determined that grazing results in increased seagrass productivity. Van Montfrans et al. (1982), Hootsmans & Vermaat
(1985) and Neckles et al. (1993) found the biomass of Zostera marina in grazed treatments was higher than in controls. Howard & Short (1986) used two large
outdoor tanks, one with grazers removed, to monitor seagrass growth. After 3 months, the above and belowground biomass
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of Halodule wrightii was significantly greater in the grazed treatments. They attributed this difference to decreased competition from epiphytes for light. Similarly,
Caine (1980) found that, when the caprellid amphipod Caprella laeviuscula was excluded from experiments, periphyton on Zostera marina increased by 411%.
Although this is a significant increase, it is not clear whether the periphyton was growing on the eelgrass or whether the figure was extrapolated from algae grown on
experimental glass slides. Unlike the laboratory studies above, we are not aware of any field studies that experimentally demonstrate that grazing increases seagrass
productivity. However, given the logistical difficulties of carrying out such an experiment in the field, this is not surprising.
Although studies of epiphyte grazing on seagrasses report a positive effect on seagrass production, studies of epiphyte grazing and fouling on seagrass are
ambiguous. Another positive effect of the removal of heavy epiphyte loads is to reduce drag and increase seagrass flexibility (Heijs 1985b,c). Removal of epiphyte
biomass by grazers may reduce fragmentation of Thalassia hemprichii leaves (Heijs 1985c).
The production of faecal material by grazers probably increases the cycling of nutrients that are then available for uptake by seagrasses and epiphytes. Robertson &
Mann (1982) suggested that intense grazing by the gastropod Littorina neglecta may be a route by which nitrogen is fixed, as the converted algal tissue becomes
available to eelgrass and other primary producers. Peduzzi (1987) suggested that molluscs have a low absorption efficiency compared to their high feeding rates, with
70% of carbon ingested by the trochid Gibbula umbilicaris, from macroepiphytes on Posidonia oceanica, being lost during defaecation. The role of bacteria in
utilizing this energy during decomposition was demonstrated by Peduzzi & Herndl (1986); a role that may be important when considering contributions to detrital
nutrient pools.
Impact of eutrophic conditions on grazerepiphyte interactions

The interaction of seagrass, grazers and epiphytes becomes crucial when attempting to predict the impact of changing nutrient levels due to anthropogenic inputs. The
link between eutrophic conditions with increased nutrient levels and the proliferation of micro and macroalgae is well established (SandJensen 1977, Cambridge &
McComb 1984, Borum 1985, Silberstein et al. 1986, Lukatelich et al. 1987, GeertzHansen et al. 1993, Neckles et al. 1993, 1994). The effect on the associated
biota of this increase in the size of the algal community is less well documented. Only a few studies have discussed a link between eutrophic conditions and changed
grazer interactions but these have not usually been tested experimentally.
Borum (1987), working on Zostera marina in Denmark, found that the eutrophic epiphyte community contributed 4 to 5 times more to the total aboveground
production in an eelgrass bed than was indicated by its biomass. The high biomass turnover was attributed to grazing. GeertzHansen et al. (1993) found that the
growth of Ulva lactuca in a eutrophic Danish estuary was controlled by isopods (Idotea spp.). In the outer parts of the estuary, where Idotea spp. were abundant,
Ulva lactuca was grazed almost to extinction, whereas in the inner parts of the estuary, where isopods were not common, large biomass accumulations of U. lactuca
occurred. In a welldesigned laboratory study, Neckles et al. (1993) studied the combined interactions of nutrients, grazers, epiphytes and seagrass productivity. By
using seasonal replicates, Neckles et al. (1993) found that the effects of nutrients and grazers on the productivity of Zostera marina varied with season. When grazer
densities were low in early summer, removing the grazers and increasing the nutrients reduced seagrass production. With high grazer
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densities in late summer, nutrient enrichment reduced seagrass production only when grazers were removed. She concluded that the influence of epiphytes on seagrass
was related to seasonally changing water temperature, grazing pressure and macrophyte requirements for light. Although Williams & Ruckelshaus (1993) did not
examine seasonal patterns, their findings also highlighted the complex nature of grazerepiphyteseagrassnutrient interactions. Their study emphasized the importance of
considering nutrient contributions from the sediment as well as the water column and that consideration of sediment nitrogen, epiphytes or grazers alone was unlikely to
result in a predictable understanding of seagrass productivity.
Conclusions
Seagrasses play a major role in providing the basic habitat for the growth of epiphytes and subsequent interactions with grazers. The number of seagrass species, range
of morphologies and environmental conditions where they are found offer very variable conditions for epiphyte colonization, growth and survival. Some seagrass
species (e.g. Thalassia and Posidonia oceanica) are similar in basic morphology and are found in similar environmental conditions. Other species, such as Zostera
marina, have different plant morphologies and life histories and occur in different conditions. The life history characteristics of a seagrass exert a significant influence on
epiphytes, which in turn interact with grazers, so care is needed when extrapolating the results from one study to another. While the underlying processes may be
similar, patterns and rates of change may be quite different, as a result of different biological and physical conditions.
Epiphytes have a wide variety of life history characteristics and morphologies that have implications for the animals that graze them. Many epiphyte species are
cosmopolitan. Some are permanent and some seasonal; the timing of their presence or absence may have a significant impact on grazer survival and abundance. The
composition, abundance and biomass of epiphytes are affected by the occurrence of freeliving reproductive stages, physical factors such as light, water movement and
temperature, as well as biological factors such as seagrass species and grazer effectiveness.
Grazers cycle nutrients in the seagrass meadows. Of the many different types of grazers, probably the most studied, and arguably the most important, are the
molluscs and crustaceans. Their diverse feeding morphologies affect their ability to graze epiphytes. Most are equipped to remove efficiently periphyton and microalgae,
including the propagules of macroepiphytes, although a few may also be adapted to grazing encrusting coralline algae and macroepiphytes. There is some evidence to
suggest that crustacean grazers can change from herbivory to detritivory.
The combination of the interactions of grazers, epiphytes and seagrasses produces the observed patterns of plant growth and abundance within the seagrass
meadow. The interactions are complex and nonlinear. They can operate on many spatial and temporal scales, ranging from very small spatial scales such as the
position on a seagrass leaf to large scales such as the variation between seagrass meadows and along environmental gradients. Temporal influences also vary in
magnitude from diel effects to seasonal and yearly cycles.
The types of interactions include direct grazing, which reduces the epiphyte biomass and removes a significant quantity of epiphyte production (between 20% and
60% of total primary production) from the seagrass meadow. The assimilation efficiencies of grazers follow a similar trend, being higher for the simpler types of
epiphytes (<70%) than the macro
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epiphytes (<20%). Grazing and food preferences may also result in changes to species composition of epiphytes. There is some evidence that grazers of seagrass
epiphytes may be limited by food supply but there is also evidence that grazer populations may be structured by predation. Although filterfeeding molluscs are
generally considered to be grazers of plankton, their role in grazing propagules of seagrass epiphytes is not known.
Other interactions between grazers and epiphytes are concerned with habitat and shelter. Epiphytes are a food source for grazers and provide habitat and shelter
from predation. The preference for habitat and the effectiveness of shelter appear to be functions of the morphology of the epiphyte and the size of the grazer. Small,
intricately branched epiphytes provide better shelter for small crustacean grazers, while larger grazers may survive better on larger, thalloidshaped epiphytes.
Grazers can significantly reduce epiphyte biomass, which results in increased aboveand belowground seagrass production. If epiphyte growth is faster than the
grazing rate, seagrasses will be overgrown and may eventually die because of shading or by being ripped from the substratum due to increased hydrodynamic drag from
epiphyte buildup. In addition, if grazing species are obligate grazers of microalgae, epiphytes may reach a sizerelated refuge from being grazed as they grow. This can
also occur with grazers that eat macroalgae if they have a preference for simpler forms and if microalgae are abundant. The ability to remove periphyton and macro
epiphytes may therefore be a function of grazing rate, food preference, growth rate of propagules and the timing and intensity of algal colonization and grazer
recruitment. Increases in nutrients may also have different effects on the seagrass community, depending upon the time of year. The recruitment dynamics of both
grazers and epiphyte propagules may therefore critically affect the outcome of these interactions (Orth 1992).
The influence of nutrients on the seagrassgrazerepiphyte community is an increasing problem throughout the world but predicting the impact of nutrients relies on a
detailed knowledge of the major processes taking place on a variety of spatial and temporal scales. The localized effect of the species, morphology and life history
characteristics of the seagrass, and the recruitment patterns of epiphytes and grazers, physical conditions and other seasonal effects such as changing temperature,
combined with the added complication of changing nutrient levels, both in the water column and sediment, mean that extrapolating the results of studies from one
location or time to another may be unwise.
Further research on the uncertainties of grazerepiphyte interactions is needed before accurate and reliable predictions of their impact on seagrass communities are
possible. This research includes better understanding of the recruitment and growth dynamics of both epiphytes and grazers and research on the dietary preferences of
grazers (including assessment of the quality of food), knowledge of local factors likely to affect interactions and also assessment of the likely magnitude of heterogeneity
at a range of spatial and temporal scales.
Acknowledgments
This review was improved by comments from G.Kendrick, H.Kirkman, N.Loneragan, T.Ward and V.Mawson. The Water Authority of Western Australia provided
partial funding for this work. Since this review was submitted there have been a number of other papers that have been published on grazerepiphyteseagrass
interactions. Unfortunately these could not be included in this review and the authors would like to apologise for their omission.
Page 153
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BEACH CLAM FISHERIES

ANTON McLACHLAN,1 JENIFER E.DUGAN,2,6 OMAR DEFEO,3 ALAN D.ANSELL,4 DAVID M.HUBBARD,2,6 EDUARDO
JARAMILLO5 & PABLO E.PENCHASZADEH7
1Zoology
Department, University of Port Elizabeth, PO Box 1600, Port Elizabeth 6000, South Africa.*
2Marine
Science Institute, University of California Santa Barbara, Santa Barbara, CA 93106, USA.
3Instituto Nacional de Pesca, Constituyente 1497, Casilla de Correo 1612, 11200 Montevideo, Uruguay.
4Scottish Association for Marine Science, Dunstaffnage Marine Laboratory, PO Box 3, Oban, Argyll, Scotland.
5Instituto Zoologia, Universidad Austral de Chile, Valdivia, Chile.
6Department of Marine Science, University of Otago, PO Box 56, Dunedin, New Zealand.
7INTECMAR, Universidad Simón Bolivar, Ap. 89000, Caracas 1080, Venezuela and Museo Argentino de Ciencias Naturales—CONICET, A.
Gallardo 470, Buenos Aires, Argentina.
* Present address: College of Science, Sultan Qaboos University, PO Box 36, AlKhod 123, Muscat, Sultanate of Oman.

Abstract The biology, ecology, and fisheries of 15 species of clam from exposed ocean beaches are reviewed and contrasted. The species,
representative of four families of bivalves, are Tivela stultorum and Siliqua patula from North America, Tivela mactroides, Donax denticulatus
and D. striatus from the Caribbean, Mesodesma mactroides and M. donacium from southern South America, Donax trunculus from Europe, D.
serra from southern Africa, D. cuneatus and D. faba from Asia, Donax deltoides from Australia, and Paphies ventricosa, P. subtriangulata and
P. donacina from New Zealand. These clams tend to fall into two categories: generally larger temperate species that dominate the macrofauna
community biomass on beaches of the dissipative type, and generally smaller tropical species, mostly donacids, found on reflective beaches. Some
species have intertidal distributions, but most are centred in the swash zone or shallow subtidal. Vertical distribution appears to be related to latitude
and temperature with lower temperatures leading to a more downshore distribution. Subtidal species are more difficult to exploit because of the
protection afforded by high energy surf zones. All are filter feeders, playing important roles in the trophic structure of beaches. Most species have
extended spawning, often with two peaks in the year. In many cases recruitment occurs in a different zone from the adult populations with subsequent
migration up or downshore. Life spans range from 1–3yr for the smaller, warm water species to >20yr in the larger temperate species, but most
species live for 2–8yr and have relatively rapid growth to maturity. Many populations exhibit resurgences—considerable fluctuations in abundance
coupled to variable recruitment and/or mass mortalities. Exploitation of beach clams is sometimes constrained by the accumulation of toxins, such as
those associated with blooms of toxic algae, that can render them unsafe for human consumption and may cause mass mortalities of the clams
themselves. Many beach clam species support recreational, artisanal and commercial fisheries, but recreational fisheries are also mostly commercial
and, with a few notable exceptions, most of the commercial fisheries are also artisanal. Recreational fisheries are notoriously difficult to manage since
numbers of harvesters cannot usually be controlled and exploitation must be limited solely by recourse to size, bag and season and/or area restrictions.
The recreational experience provided by clam fisheries on ocean beaches must be considered to be as valuable as the food value of the resource itself.
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Introduction
Exposed sandy beaches dominate the icefree shores of all continents and provide important recreational and commercial resources in many areas. Because the fauna
of ocean beaches is cryptic and mobile it is easily overlooked and has tended to be neglected by ecologists. Nevertheless, many beaches, especially those of the
dissipative type (Short & Wright 1983), support rich biotas of which clams are often an important component, especially in terms of biomass. On all continents at least
one clam from exposed beaches is extensively collected as part of a recreational, artisanal or commercial fishery (Fig. 1). Large species from temperate areas are most
often targeted, but there are exceptions.
For the purposes of this review we define a recreational fishery as collection for bait or food without sale or dependence on the resource; an artisanal fishery as
collection for subsistence or sale by individuals or groups using traditional methods; and a commercial fishery as collection for sale by corporate or collective
organizations. However, these distinctions are not precise; in many cases artisanal fisheries are also commercial (e.g. Chile, Uruguay) and recreational fisheries often
also support commercial activities. Clam fishery catches, where available, are expressed as recorded landings. These are not generally based on stock size and they
reflect a variety of influences, such as changes in market, economic and political climates, regulations, etc., and so must be interpreted with care.
The aim of this review is to summarize available information on beach clam fisheries, to contrast patterns in the biology, ecology and fisheries of the species involved
in relation to beach type, geographical area and socioeconomic factors, and to assess the current status of clam fishery management on beaches that, in most cases, are
also used for other recreational purposes. Fifteen species from exposed ocean beaches and surf zones are covered (Table 1, Fig. 2). All are heterodont
eulamellibranchs except Siliqua, which is an adapedont. Not all potentially exploitable beach clams are included, only those species for which sufficient information is
available to allow meaningful comparison. Even so, the 15 species have not been equally well studied; in some cases poorly researched species are included for
comparison and because they are considered important in their region.
Figure 1 Geographical distribution of the species of beach clams considered in this review. North America: a, Tivela stultorum; b,
Siliqua patula; Caribbean: c, Tivela mactroides; d, Donax denticulatus; e, D. striatus. South America: f, Mesodesma
mactroides; g, M. donacium, Europe: h, Donax trunculus. Africa: i, D. serra; Asia: j, D. cuneatus; k, D. faba. Australia: l,
D. deltoides. New Zealand: m, Paphies ventricosa; n, P. subtriangulata; o, P. donacina.
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Table 1 Taxonomy and common names of the species dealt with in the text.
Geographical area Species Authority Superfamily Family Common name

North America Tivela stultorum (Mawe) Veneracea Veneridae Pismo clam
Siliqua patula (Dixon) Solenacea Solenidae Pacific razor clam
Caribbean Tivela mactroides (Born) Veneracea Veneridae guacuco (Venezuela);
Donax denticulatus (Linnaeus) Tellinacea Donacidae beach clam (Jamaica)
Donax striatus Linnaeus Tellinacea Donacidae chipichipi (Venezuela)
South America Mesodesma mactroides (Deshayes) Mactracea Mesodesmatidae yellow clam
Mesodesma donacium (Lamarck) Mactracea Mesodesmatidae macha
Europe Donax trunculus (Linnaeus) Tellinacea Donacidae wedge clam
Africa Donax serra (Roding) Tellinacea Donacidae white sand mussel
Asia Donax cuneatus (Linnaeus) Tellinacea Donacidae
Donax faba (Gmelin) Tellinacea Donacidae
Australia Donax deltoides Lamarck Tellinacea Donacidae pipi, Goolwa cockle
New Zealand Paphies ventricosa (Gray) Mactracea Mesodesmatidae toheroa
Paphies subtriangulata Wood Mactracea Mesodesmatidae tuatua
Paphies donacina (Spengler) Mactracea Mesodesmatidae tuatua
North America
Two species of beachdwelling bivalves are harvested on beaches of the northeastern Pacific coast. These were among the largest beach clam fisheries in the world in
terms of yield and also the most extensive in terms of shoreline length and latitude. No other beach clams are collected on any comparable scale in the USA.
The Pismo clam, Tivela stultorum (Fig. 2a)
Distribution
Tivela stultorum occurs on the Pacific coast of North America from Half Moon Bay, California to Isla Socorro in the Islas Revillagigedos, Mexico (Fitch 1950) (Fig.
1). Pismo clams can be rare to common on beaches in this range. The northern limit of the distribution of T. stultorum varies, expanding and contracting over time with
episodic settlement events (Coe 1953). Populations of T. stultorum occur in a variety of sandy habitats including high intermediate to dissipative sandy beaches and the
sandy entrance channels of bays, sloughs and estuaries (Shaw & Hassler 1989). T. stultorum can be found from the lowermid to low intertidal and subtidally to
depths of 25m (Shaw & Hassler 1989). The species is particularly prevalent in upwelling areas (Coe & Fitch 1950). In Monterey Bay, clams occurred on beaches
with slopes of 1.1–4.6 degrees and not on those with steeper slopes (Nybakken & Stephenson 1975).
Biology and ecological role

T. stultorum is a broadcast spawner and fertilization is external (Coe 1947). The sexes are separate and similar in appearance, and hermaphrodites are rare (<1%)
(Nybakken & Stephenson 1975). Males usually spawn first and the release of male gametes stimulates female spawning (Shaw & Hassler 1989). The reproductive
cycle of T. stultorum is seasonal in Californian waters (Coe & Fitch 1950) and multiple spawnings may occur during a
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Figure 2 The species of beach clams covered in this review; key to species as in Figure 1.
season. Gametogenesis commences in March or April and ripe gametes are first evident in April and May. Spawning begins in late July or early August and continues
through November (Coe 1947). Peak spawning occurs during September and October in Californian populations (Coe 1947, Nybakken & Stephenson 1975). The
spawning season is extended in populations in Baja California (Coe & Fitch 1950). Fecundity increases with female size. In clams of legal size, 114mm or larger,
fecundity ranges 11–20 million eggs and averages 15 million eggs per season (Coe 1953, Fitch 1950, 1961). The eggs are 74 to 81 μm diameter (Coe & Fitch 1950).
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The larval biology of T. stultorum is not well known. Larval development is estimated to take 3 weeks (Coe 1947). Larvae reared in the laboratory are negatively
buoyant (A.Hazeltine pers. comm) and it is suggested that they spend much of their larval life near the bottom, moving little in surface currents (Shaw & Hassler 1989).
At metamorphosis and settlement the clam develops a foot and attaches to sand grains with a byssus (Shaw & Hassler 1989). As the clam grows the byssus
disappears. Young clams occur in the mid to low intertidal zone of beaches. Settlement of T. stultorum has been observed in the intertidal, but it is not known whether
settlement also occurs subtidally.
Pismo clams are suspension feeders, removing phytoplankton and detritus from the overlying water (Coe 1947). Because they have short siphons, feeding and
undisturbed animals are buried close to the sand surface. The inhalant siphon has elaborate frills to reduce sand ingestion but the gut is often 50% full of sand (Fitch
1950).
T. stultorum is a bulky bivalve (114mm length, minimum legal size, weighs 350g live) with a large foot. Adult clams are not capable of much movement on the
surface of the sand and have not been observed to migrate tidally (Fitch 1950), although small clams may be more mobile than large ones. The heavy shell appears to
assist in burrowing and maintaining position in the sediments. Adult clams burrow to depths of 50–160 mm (Shaw & Hassler 1989). This species has been described
as a slow burrower (Shaw & Hassler 1989) but, in a comparison of rates of burrowing in sandybeach bivalves, the rate for T. stultorum was found to be intermediate
relative to other species (McLachlan et al. 1995). A study of T. stultorum outplanted to intertidal and subtidal locations found animals planted subtidally occurred in
subsequent intertidal surveys, indicating that subtidal animals moved into the intertidal zone (Dugan et al. 1995).
The shells of T. stultorum often carry a commensal hydroid, Clytia bakeri, and fishers may use the hydroid tufts to locate clams (Fitch 1950). A commensal pea
crab, Pinnixia sp., occurs occasionally in the mantle cavity (Frey 1971). A larval cestode that matures in stingrays and skates occurs in some clams and stops the
sexual development of infected individuals (Warner & Katkansky 1969). Trematodes have been reported in some populations of clams (Coe & Fitch 1950). No
diseases have been described in T. stultorum and incidents of mass mortalities are rare (Fitch 1950).
T. stultorum plays an important role in the filtering of large quantitities of sea water and the conversion of surf zone primary production to secondary production. It is
the dominant macrofauna species on some beaches, with high abundances and densities (>980 mature clams m−1, Fitch 1950).
T. stultorum is an important prey item for a number of vertebrate and invertebrate predators and experiences considerable natural mortality at all ages. Skates, rays,
surf fish, rock crabs, moon snails, gulls and sea otters prey upon T. stultorum (Miller et al. 1975, Haderlie & Abbott 1980). Most predation occurs at high tide with
the exception of some of the gull predation. Moon snails, Polinices lewisi, drill the shells and prey on clams of up to 2yr (42–50mm) in age (Fitch 1950, Shaw &
Hassler 1989). Rock crabs, Cancer sp., prey on clams up to 25mm in length by cracking the shell (Fitch 1950, Shaw & Hassler 1989). Gull predation on T.
stultorum can be enhanced by the activities of human fishers that strand clams on the surface (Fitch 1950).
Some studies have implicated sea otter (Enhydra lutris) predation in recent (early 1970s) declines of pismo clam populations and fisheries in California (Stephenson
1977, Wendell et al. 1986). A sea otter was observed to eat 24 clams in 2.5h (Stephenson 1977). Wendell et al. (1986) estimated that one sea otter can consume 80
clams per day. In Monterey Bay, Miller et al. (1975) estimated that sea otters consumed 520000 to 700000 clams in a year.
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Sea otters appear to feed primarily on clams that are 60mm in length or larger (Estes & Van Blaricom 1985). The average size of clam consumed by sea otters in
Monterey Bay was 127mm (Nybakken & Stephenson 1975). Estes & Van Blaricom (1985) question the relative effects of sea otters on Tivela stultorum in the light
of other influences on clam populations, including disturbance of clam beds by shellfish fishers, poaching, heavy motor vehicle traffic on clam bearing beaches, and the
naturally high mortality of juvenile clams on certain beaches.
Population dynamics
Populations of T. stultorum have been extremely variable in time and space and were classed with those of Donax gouldi and other bivalves as resurgent by Coe
(1953). This parallels the observation that settlement and recruitment of Tivela stultorum is extremely variable from year to year (Coe 1953). Existing data suggest
that recruitment is independent of stock size (Tomlinson 1968). Detectable settlement events may occur once a decade or so and major settlement events occur
episodically, at 18–28yr intervals, on California beaches (Estes & Van Blaricom 1985, Shaw & Hassler 1989). Recruitment of T. stultorum also varies significantly
between beaches in any one region, e.g. Pismo Beach and Astascadero beach in the early 1970s (Wendell et al. 1986). As a result of the infrequent recruitment
success and the long life of this species, populations of T. stultorum are often skewed towards large old animals. Fitch (1965) described a subtidal population where
all but one clam was older than 13yr and only 2.5% were smaller than 118mm (n=200).
Tivela stultorum is a longlived species, often living beyond 20yr and attaining sizes of 150mm or greater (Fitch 1950) (Table 2). The largest clam reported was
187mm long, weighed 1589g and was estimated to be 26yr old (Carriquiriborde et al. 1981). Gillilan (1964) estimated the age of one individual from Zuma beach,
California to be 53yr. For the first 2 yr on the beach, pismo clams grow 21–25mmyr−1 (Coe & Fitch 1950). Growth rates decline with increasing age and 3 to 6yr
old clams grow at rates of 10–18 mmyr−1. Data
Table 2 Summary of key features of beach clam fisheries reviewed in the text. Under habitat I=intertidal, S= subtidal. Under fishery type A=artisanal,
C=commercial, R=recreational and *=no longer operational. Lengths in mm and landings in tonnes live weight per year. =log10(K)+ 2log10
(L∞ ) (approximate average values for each species are presented).
Species Habitat Length at maturity Lifespan Legal size Max length Fishery type Maximum landings
Donax deltoides I&S 36 3.5 none/35 60 C&R unknown 3.70
Mesodesma mactroides I 43 3.5 50 85 A, C & R 1078 (1953) 3.71
Mesodesma donacium S 50 15 50–60 100 A&C 18000 (1989) 3.74
Tivela stultorum I&S 25 >20 114–127 187 R&C 24000(1945) 3.71
Siliqua patula I&S 100 5–19 85–114 170 C&R 3600 (1915) 4.02
Donax serra I&S 44 >5 58 80 R & C* 20 (1968) 3.57
Donax trunculus I&S 8 1.5–>6 30 (Spain) 44 R&A unknown 3.08
Tivela mactroides I&S 20 1.5 – 35 C&R 354 (1992) 2.33
Donax denticulatus I 10 1.5 – 23 A 226 (1990) 3.24
Donax striatus I 10 1.5 – 35 A unknown 3.15
Donax cuneatus I 12 <3 – 23 A unknown 2.55
Donax faba I 10 <3 – 27 A unknown 2.64
Paphies ventricosa I 40 >10 100 160 R* & C* 185 (1940) 3.83
Paphies subtriangulata I&S unknown >5 none 85 R&C 131 (1993) 3.30
Paphies donacina I&S unknown 17 none 109 R&C 5 (1992) 3.47
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from Fitch (1950) allow an estimate to be made of the growth rate index values and pay less attention to individual values of K and L∞ . Legal harvest size (114mm
or 127mm) for T. stultorum is reached in 5–8yr in California (Coe & Fitch 1950).
Growth of T. stultorum varies seasonally and geographically, and slows markedly during the winter months (Coe & Fitch 1950) in California populations. However,
slower winter growth was not observed in T. stultorum inhabiting beaches in Baja California (Searcy Bernal & Saavedra Rosas 1988). In general, faster growth of T.
stultorum is observed in southern California populations than in northern California populations (Coe & Fitch 1950). Within a region, growth rate of clams can vary
from beach to beach (Herrington 1929, Coe 1947, Coe & Fitch 1950). Pismo clams mature at shell lengths of 17–30mm (Coe 1947). Sexual maturation occurs during
the first full summer in populations on southern California beaches but only during the second summer in populations near the northern limit of its range (Shaw & Hassler
1989).
The age and growth rates of T. stultorum are usually determined by examination of annuli or growth rings on the shell (Fitch 1950, Searcy Bernal et al. 1989, Shaw
& Hassler 1989). These rings are usually formed in the fall or winter months in California and can be seen externally and internally in crosssections of valves. External
annuli are more easily distinguished on clams less than 5yrold and become less distinct as growth slows (Fitch 1950). Searcy Bernal et al. (1989) suggest that analyses
combining internal and external rings result in the most accurate age and growth rate determination in this species.
Sources of mortality other than predation for T. stultorum include natural events, fishing and pollution. Storm surf events can cause significant mortality in clam
populations (Fitch 1950). Tomlinson (1968) estimated fishing mortality of T. stultorum of all ages as 0.8yr−1. Occasional large mortalities, particularly of juvenile and
young clams, have been attributed to heavy freshwater runoff, extreme cold and petroleum oiling (Fitch 1950).
Fishery
T. stultorum has been consumed and used by humans for thousands of years. The clams occur in middens of the coastal Chumash Indians, who used the shells as
ornaments and tools (Frey 1971). Pismo clams are highly prized as delicacies in California but are not for sale in markets. In the USA, products of the now closed
commercial fishery included fresh clams, canned clam meat and livestock food. The USA commercial fishery for T. stultorum lasted from 1916 to 1947 and occurred
primarily at California beaches near Morro Bay, and at Pismo Beach, Oceano and Monterey Bay (Shaw & Hassler 1989). USA landings peaked in 1918 (302t) and
declined thereafter despite management and restoration efforts (Fig. 3). Pismo clams were one of the most important commercial littoral invertebrate species in Baja
California until the early 1980s (Searcy Bernal & Saavedra Rosas 1988). Between 1935 and 1947 up to 24000tyr−1 were imported to the USA from Mexico for
canning to supplement declining California landings (Shaw & Hassler 1989). Shipping was discontinued in 1947
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Figure 3 Commercial fishery landings of Tivela stultorum in California from 1916–47. (After Shaw & Hassler 1989).
due to high costs and the closure of the California fishery. Landings of T. stultorum in Baja California from 1977 and 1978 exceeded 1000tyr−1 (Searcy Bernal 1983)
and commercial harvesting continues in Baja California, Mexico, primarily at beaches near San Quintin.
The extent of recreational harvest of T. stultorum in Mexico is unknown. Recreational harvest occurs throughout the USA range of the species and affects both
intertidal and subtidal populations. The total recreational landings for California are not known but thousands of people participate in clamming. Landings are variable
and have declined in recent years despite some significant recruitment events. For example, in 1972 the largest settlement of T. stultorum in 30yr was observed at
Atascadero Beach, California (Wendell et al. 1986) and landings did not increase.
To illustrate the pattern seen in many areas, recreational landings for Pismo Beach for 1975–83 are given in Figure 4. The decline in landings at Pismo Beach and
other northern clamming areas seen in the 1970s and early 1980s were attributed to the expansion of the sea otter’s range by some investigators (Miller et al. 1975,
Stephenson 1977, Wendell et al. 1986). Southern California clam landings also declined precipitously in the early 1980s, this being attributed to catastrophic storms
associated with the 1982–83 El Niño Southern Oscillation (ENSO) event (P.Haaker, California Dept of Fish and Game, pers. comm.). A series of good recruitment
years in the late 1980s have revived the collapsed recreational fishery for T. stultorum on central and southern California beaches through the early 1990s (J.E. Dugan
pers. obs.).
The failure of the commercial harvest in the USA and the high variability of the recreational harvest attest to the difficulty of sustainable management of T. stultorum
fisheries with conventional approaches. Harvests of T. stultorum are apparently based on the strength of occasional, unpredictable strong year classes. Once a strong
cohort or group of cohorts is fished down or affected by natural events or catastrophes, the harvest collapses until the next period of strong recruitment. Recreational
landings from Pismo Beach vary considerably over time but illustrate that point (Fig. 4).
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Figure 4 Recreational fishery landings of Tivela stultorum at Pismo Beach 1975–83. (After Wendell et al. 1986).
In California, early efforts to manage the fishery included bag limits, size limits, area closures and transplants of brood stock (Fitch 1950). The bag limit was reduced
from 200 clams in 1911 to 50 clams in 1915, to 10 clams in 1948 (Fitch 1950). The minimum legal size is smaller in southern California (114mm for San Luis Obispo
County and south) than in northern California (127mm for Monterey County and north) (Shaw & Hassler 1989) (Table 2, p. 168). Fishing gear is restricted to hand
tools, e.g. a sixpronged potato fork, rakes, hands and feet. Divers locate siphons then excavate subtidal clams. None of these regulations have restored the T.
stultorum fisheries to anything resembling the pre1920 levels nor have they resulted in sustainable harvests.
The difficulty of controlling recreational harvest effort is well illustrated by the T. stultorum fishery in California. Anyone can purchase a clam fishing licence;
therefore only season and area closures, and bag and size limits can be used to manage the fishery. Recreational fishing pressure can be great; for example, over
4million clams were removed in a 10wk period from a 6.4km stretch of beach (Frey 1971). On one weekend on Pismo Beach, an estimated 150000 clam diggers
visited the beach and removed 341 of clams (Fitch 1950). Incidental mortality of undersized clams, which are often discarded on the surface and fall prey to gulls,
desiccation and thermal stress, is a problem in this fishery (Shaw & Hassler 1989).
Although pismo clams can accumulate PSP (paralytic shellfish poisoning) toxins produced by blooms of certain dinoflagellates, no cases of human PSP poisoning
were reported during the height of the fishery (Fitch 1950). Seasonal shellfish closures, known as quarantines, are imposed throughout California to prevent any human
consumption of toxic shellfish during the summer season when toxinproducing phytoplankton blooms are known to occur.
A traditional fishery model (yield per recruit) developed by Tomlinson (1968) using unfished populations, suggested that the survival of T. stultorum increased with
age. The current minimum size limit of 114mm in Southern California compared well with age at entry for maximum yield. Recruitment was independent of total
mortalities in this model, which used an instantaneous fishing mortality of 0.8yr−1 for all ages but recognized that it was possibly an underestimate of present fishing
mortality.
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The Pacific razor clam, Siliqua patula (Fig. 2b p. 166)
Distribution
S. patula occurs on the eastern Pacific coast of North America from Pismo Beach, California to the Aleutian Islands, Alaska (Lassuy & Simons 1989) (Fig. 1, p.
164). In British Columbia the clams are abundant only at Long Beach on the west coast of Vancouver Island, and near Masset on the northeastern coast of Graham
Island (Queen Charlotte Islands). Populations of S. patula inhabit stable, open coast sand beaches, occurring primarily on fully exposed dissipative type beaches that
are broad, flat, hard and composed of fine sand (Lassuy & Simons 1989). Large adult clams are densest in the low intertidal zone but subtidal populations can be
substantial in some locations. Juvenile clams occur both intertidally and subtidally (Lassuy & Simons 1989). Adults are usually buried in the sand at depths of 300mm or
more (McMillin 1924) and juveniles at about 50mm (Lassuy & Simons 1989).

S. patula is a broadcast spawner and fertilization is external. The sexes are separate and the sex ratio of adult clams is 1:1 (Nickerson 1975). The eggs are c. 90μm in
diameter (Breese & Robinson 1981) and are negatively buoyant (Weymouth et al. 1925). The peak spawning period varies geographically: in the late spring or early
summer in Oregon and Washington (Lassuy & Simons 1989), and in August in Alaska (Weymouth et al. 1925). Some spawning may occur yearround and be related
to food availability (Lassuy & Simons 1989). The initiation of spawning in S. patula may be related to water temperature, a rapid increase in water temperature, food
availability, upwelling, the tidal cycle or some combination of those factors (e.g. Weymouth et al. 1925, Bourne & Quayle 1970, Nickerson 1975, Breese & Robinson
1981, Lassuy & Simons 1989).
The larval development of S. patula spans 5–16 wk at temperatures of 11–15°C, with the following stages: appearance of a veliger at 10 days, metamorphosis to a
clamlike shape after 3wk, a distinct foot at 5wk, at 8wk loss of the velum and the shell becoming opaque, followed by settlement after 10wk (McMillin 1924). It has
been suggested that razor clam larvae are negatively buoyant and spend much of their time in sand, thus limiting larval dispersal (McMillin 1924, Weymouth et al.
1925). However, recent observations indicate that the larvae are not negatively buoyant and do not occur primarily in the sand (D.Simons, pers. comm.). Settlement of
S. patula can apparently occur intertidally and subtidally (Lassuy & Simons 1989). Large numbers of juvenile clams have been observed in both habitats (McMillin
1924, Rickard et al. 1986).
Adult razor clams are rapid burrowers, capable of vertical rates of movement of 230–305mm min−1 (McMillin 1924, Lassuy & Simons 1989). S. patula displayed
the highest burrowing rate index values of 12 species investigated by McLachlan et al. (1995). Adult razor clams show little lateral movement but juvenile clams are
capable of some lateral movement (Bourne & Quayle 1970). The southern limits of S. patula distribution may be related to water temperatures and climate. In the
laboratory, lethal temperatures for razor clams in sea water ranged from 27.5–22.5°C for exposure times of 1–4h, respectively (Sayce & Tufts (1971).
A commensal nemertean, Malacobdella grossa, 25–50mm in length, occurs inside the mantle cavity and siphon of Siliqua patula and a commensal pea crab,
Pinnixia sp., is regularly found in the mantle cavity (Lassuy & Simons 1989). An infectious gill disease appeared in Siliqua patula populations in 1983, causing major
mortalities and resulting in the closure
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of the Washington fishery in 1984 and 1985 (Lassuy & Simons 1989). The disease was identified as nuclear inclusion X (NIX) by Elston (1986) and was unknown
prior to that outbreak. It is caused by a prokaryotic, intranuclear bacterium that affects the gills, resulting in serious tissue damage to the gill epithelia and secondary
bacterial infections (Elston 1986, Ay res & Simons 1994).
Pacific razor clams are filter feeders and can form a dominant component of the sandbeach community. Surfzone phytoplankton, particularly the principal diatom
involved in surfzone blooms, Chaetoceros armatum, probably compose the bulk of the diet of Siliqua patula (Lewin et al. 1979a). Lewin et al. (1979b) suggest that
dense populations of razor clams may play an important role in surfzone nitrogen cycles through the excretion of ammonium. Juveniles and new recruits are consumed
by birds (sandpipers, glaucouswinged gulls, surf scoters, ducks and crows), fish (English sole recruits, sturgeon and surf perches) and young Dungeness crabs
(Cancer magister) (McMillin 1924, Tegelberg & Magoon 1969, Hogue & Carey 1982, Lassuy & Simons 1989). Sea otters, Enhydra lutris, prey upon adult clams
throughout their range, particularly in Alaska (Estes & Van Blaricom 1985).
Population dynamics
Recruitment variability in populations of Siliqua patula may be related to currents and climatic events (Lassuy & Simons 1989). Populations of S. patula are
somewhat resurgent (sensu Coe 1953) in the southernmost portion of the species range as a result of the high recruitment variability in that region. Juvenile clams have
been observed to reach high densities (>16000m−2) in the intertidal zone but generally occur in densities of 0–1076m−2 in the lower third of the intertidal zone (Lassuy
& Simons 1989). A subtidal density of juvenile (1–15mm) razor clams of 38000m−2 was estimated in one study (Rickard et al. 1986).
Some workers have estimated the ages and growth rates of S. patula by counting the number of growth rings on the shell (Lassuy & Simons 1989). Each growth or
check ring represents an annual ring or annulus, although the presence of other rings or checks formed during spawning or disturbance has also been noted (McMillin
1924, Weymouth & McMillin 1931, Hirschhorn 1962). The appearance and spacing of annuli can vary geographically in S. patula. Annuli in populations of razor
clams in Alaska were more distinct, numerous and closely spaced, relative to those in Washington populations (McMillin 1924, Weymouth et al. 1925). Tegelberg
(1964) suggested that the formation of distinct annuli in S. patula is dependent upon a period of slow growth in the winter, a phenomenon that may not occur in some
populations. Growth rings or annuli are not considered reliable for use in age determination in razor clam populations in Washington State (D.Ayres pers. comm.).
S. patula is relatively long lived and has indeterminate growth. The growth rate, maximum length (120–170mm), maximum age (5–19yr), and population structure of
S. patula vary geographically and with fishing pressure (Lassuy & Simons 1989) (Table 2, see p. 168). Growth rates are higher and maximum lengths, size at maturity
and life spans are lower in southern populations (Lassuy & Simons 1989). For example, at Pismo Beach, California, at their southern distribution limit, razor clams
grow to 127mm, live up to 5yr and have growth rates is lower (3.78) than in the southern population.
The growth of S. patula varies with tidal height; higher growth rates occur in clams living
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near the low tide line and lower growth is found in those living higher on the beach (Tegelberg 1964, Bourne & Quayle 1970, Quayle & Bourne 1972). Growth rates
of S. patula also vary seasonally; growth is usually slowest in the late fall and winter months and increases with the warmer water of spring (Weymouth et al. 1925,
Hirschhorn 1962, Tegelberg 1964).
The age at maturity of S. patula varies geographically (Weymouth et al. 1925), ranging from 2–4yr and increasing with increasing latitude (Weymouth et al. 1931).
At the southern end of the species’ range, Pismo Beach, California, clams mature at 2yr. In Alaska, near the northern end of the species’ range, clams mature at 3–4yr.
In general, sexual maturity occurs at a shell lengths of c. 100mm (Lassuy & Simons 1989) (Table 2, p. 168).
Natural mortality of juvenile razor clams from predation by marine and terrestrial predators such as birds, fish, and crabs is high. Other significant sources of mortality
to S. patula, especially younger clams, may include the scouring effects of winter storms, vehicle traffic on beaches, flood events and siltation (McMillin 1924,
Tegelberg & Magoon 1969, Bourne & Quayle 1970, Nickerson 1975, Lassuy & Simons 1989). Natural mortality of adult clams is probably lower than that of
juveniles. Sea otters prey on adult clams but their impact on populations has not been documented as it has on populations of Tivela stultorum in California (Wendell
et al. 1986). McMillin (1924) estimated that the mortality of Siliquapatula was up to 99% in the first 8 months. Annual survival estimates increased with age, 9% for
years one and two, 30% for years two and three, and 40% at greater ages (Nickerson 1975). Link (1980) found the opposite trend in his tagging study: highest
survival (15.5%) occurred at age 0 with lowest survival (0.1%) in clams older than 3yr, but he had low returns of the latter.
Fishing is a significant source of mortality in populations of S. patula. Instantaneous mortality rates (Z) of S. patula in Oregon were estimated as 2.52yr−1 and
2.34yr−1 by Hirschhorn (1962) and Link (1980). Fishing mortality (F) was estimated as 1.78yr−1 and natural mortality (M), including wastage, as 0.74yr−1 by
Hirschhorn (1962). Nickerson (1975) estimated M (not including wastage) as 0.33–0.50yr−1 for Alaskan populations of razor clams. Wastage of undersized clams is a
significant source of fisheryassociated mortality in S. patula populations (Lassuy & Simons 1989). Early studies estimated wastage to be equal to landings (McMillin
1924).
Epidemic levels of NIX disease resulted in significant mortality in S. patula populations, especially in Washington, in the early 1980s (Elston 1986). Disease
associated mortality rates of S. patula were dependent upon the prevalence and the intensity of NIX infection. The prevalence and the intensity of NIX infection in
razor clams varied geographically and temporally, as did the mortality rates. Razor clam populations experienced 95% mortality in 1983–84 along the central coast of
Washington where the prevalence of NIX was high (100% between June 1983 and June 1985) (Elston 1986). To the north and south of central Washington, the
prevalence and intensity of NIX decreased. NIX was not observed in razor clams at the Queen Charlotte Islands or further north. A high prevalence of NIX was
observed in razor clams on some Oregon beaches but the intensity of the infections was low and mortalities were not significant (Link 1986). LeClair & Phelps (1994)
suggest that variation in the susceptibility of different populations to NIX may have a genetic basis. In Washington during 1987–93, NIX was widespread in razor
clams, prevalence was high (c. 100%), intensity ranged from 2% to >40%, averaging about 20%, but no major mortality was detected (Ayres & Simons 1988, 1989,
1992). The intensities of NIX infection have generally increased over the last 10yr (Ayres & Simons 1994).
Page 175
Fishery
Razor clams were harvested by native Americans and the shells are known from middens (Lassuy & Simons 1989). The modern fishery was initiated as early as 1894
in Oregon and spread north to Alaska (Nickerson 1975). By 1915, up to 3630tyr−1 were harvested commercially in Washington alone (Lassuy & Simons 1989). Size
and season restrictions were imposed first on commercial harvest, recreational harvest being unrestricted until the 1920s, when restrictions on size, bag limits, and
season were first imposed (Lassuy & Simons 1989). The fishery has shifted from commercial to recreational in Washington and Oregon and the recreational harvest
now far exceeds the commercial harvest in the Pacific Northwest (Lassuy & Simons 1989). Up to 1940, 80% of Oregon take was commercial (Link 1980); after that
time the recreational take increased. Commercial harvests of S. patula occur in Alaska, British Columbia, Washington and Oregon (Fig. 5) but not in California
(Schink et al. 1983). Traditional fishery models for S. patula are not available in the literature.
In Alaska, commercial landings of razor clams exceeded 2200t in 1917, but have fluctuated considerably in subsequent years (Schink et al. 1983, Estes & Van
Blaricom 1985). From 1960 to 1974, landings declined, ranging from a high of 210t in 1960 to a low of 5t in 1966 (Fig. 5). Annual landings averaged 134t for 1960–
63, but after the earthquake in 1964, annual landings averaged 23t for the period through 1974 (Estes & Van Blaricom 1985). In recent years, most of the commercial
harvest of razor clams comes from scattered locations on the Alaska Peninsula, Cook Inlet, Prince William Sound, Kodiak and the Copper River Delta. That harvest is
used primarily as bait for the Dungeness crab fishery. The razor clam resources of Alaska are not fully utilized because of the distance from markets, supply fluctuations
and shellfish toxin concerns.
Figure 5 Commercial fishery landings of Siliqua patula in Alaska (1916–80), British Columbia (1981–92), Washington (1935–93) and
Oregon (1942–86) (After Schink et al. 1983, Estes & Van Blaricom 1985, Lassuy & Simons 1989, Ayres & Simons 1992,
1994, Bourne pers. comm.). Landings for Alaska from 1975–80 are for Kodiak and Prince William Sound only. Landings for
Washington from 1980–93 do not include the Quinault Indian Reservation.
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In British Columbia, the commercial harvest of S. patula occurs only at Massett in the Queen Charlotte Islands (Estes & Van Blaricom 1985). Commercial harvest
of razor clams began in 1924, producing canned clams, but the canneries shut down in the late 1960s and the commercial fishery harvest is now primarily used for
Dungeness crab bait. Since 1970, commercial landings have ranged from 18–155tyr−1 (N.Bourne pers. comm., Estes & Van Blaricom 1985) (Fig. 5).
In Washington, commercial harvest of S. patula has fluctuated dramatically since 1915 and declined in recent years (Fig. 5). The commercial fishery is now relatively
small and limited to beaches in the Willapa Spits in Willapa Bay and on the Quinault Indian Reservation (Schink et al. 1983, Lassuy & Simons 1989). This is used
primarily for Dungeness crab bait although some is sold for human consumption. Between 1950 and 1979 annual landings ranged from 77 to 467t (Schink et al. 1983,
Lassuy & Simons 1989). Since 1980, offreservation annual landings have not exceeded 35t and averaged less than 10t when the beaches were open for commercial
harvesting (harvest figures from the Quinault reservation are not available for that period). The annual value of the harvest to fishers has ranged from US$4000 to
US$89000 since 1980 (Lassuy & Simons 1989). In Oregon, limited commercial harvest primarily occurs on beaches between the Tillamook Head and the Columbia
River (Estes & Van Blaricom 1985). Most of the commercial harvest is marketed as fresh meat for human consumption. Annual commercial landings have declined
since the mid 1950s and during recent years have fluctuated considerably, ranging from 45–55t (Estes & Van Blaricom 1985, Lassuy & Simons 1989). The mean
annual landings for 1977–86 were 12t (142000 clams) with a mean total value of US$35300. The highest harvest of that period occurred in 1985 and was 26t
(303000 clams) with a value of US$115000. The lowest harvest in that period occurred in 1983 and was 45kg (1000 clams) with a value of US$100 (Lassuy &
Simons 1989).
The market price of razor clams is generally high and stable (Lassuy & Simons 1989). Retail value of razor clams range from US$2–5kg−1 unshucked and up to
US$14kg−1 shucked (Lassuy & Simons 1989). The wholesale value of commercial landings varies considerably between years and locations, ranging from US$2–
4.55kg−1 (Lassuy & Simons 1989).
Recreational harvest of S. patula occurs throughout the species range but the intensity varies. Recreational harvest landings of S. patula are greatest on the
Washington coast (Fig. 6). The recreational value of razor clams is high and the open season draws many participants, both resident and nonresident to clamming
beaches (Lassuy & Simons 1989). Recreational harvest of razor clams may have considerable indirect value to coastal communities including increased revenue from
housing and feeding of clammers during the season. Important sport harvest of razor clams occurs in the Clam Gulch area of the Kenai Peninsula and in Prince William
sound in Alaska (Schink et al. 1983, Estes & Van Blaricom 1985). Annual landings are relatively low and did not exceed 900000 clams between 1969 and 1975
(Schink et al. 1983) (Fig. 6). Levels of recreational harvest are poorly documented in British Columbia. The major recreational harvest is at Long Beach on Vancouver
Island.
The most intensive recreational harvest of S. patula occurs in Washington and landings fluctuate considerably. Between 1949 and 1976, landings ranged from 1.5–
3.5 million clams (Ayres & Simons 1992). Between 1977–86, the average sport harvest was 6.4 million clams per year, the highest harvest during that period was 13
million clams, c.585t (1979) and the lowest when the fishery was open was 3.2 million clams (1986) (Lassuy & Simons 1989). The fishery was closed in 1984 and
1985 due to NIX disease. The fishery has recovered since then and, despite closure due to high levels of shellfish toxins, annual landings have averaged over 2.7 million
clams since 1986 (Ayres & Simons 1988, 1989, 1992, 1994) (Fig. 6). Land
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Figure 6 Recreational fishery landings of Siliqua patula in Alaska (1969–75), Washington (1949–93) and Oregon (1955–85). (After
Schink et al. 1983, Lassuy & Simons 1989, Ayres & Simons 1992, 1994, Ayres pers. comm.).
ings for 1994 exceeded 4 million clams (D.L.Ayres pers. comm.).

A less intensive recreational harvest of Pacific razor clams occurs in Oregon where annual landings average less than 2 million clams (Schink et al. 1983) (Fig. 6).
Recreational harvest between 1977–86 averaged 683000 clams per year, the highest harvest during that period being 1131000 (1985), the lowest 117000 (1982–83
ENSO event) (Lassuy & Simons 1989). In California recreational harvest of razor clams occurs primarily in the northern part of the state, between Humboldt Bay and
the Smith River (Estes & Van Blaricom 1985). Recreational landings are small and the fishery is considered to be fairly stable (Schink et al. 1983).
Regulations concerning the harvest of S. patula vary between regions and between commercial and recreational harvest. In Alaska the commercial size limit is
114mm (Schink et al. 1983). In British Columbia the commercial fishery has a size limit of 85mm and no bag limit while the recreational fishery has no size limit and a
bag limit of 12–75 clams (Bourne et al. 1987). The small recreational fisheries are not closely regulated in Alaska or British Columbia. In Oregon the commercial size
limit is 95mm and seasonal closures are enforced (Schink et al. 1983) (Table 2, p. 168). The recreational fisheries in California and Oregon are controlled by
regulations on bag limits, seasons and areas.
Both commercial and recreational seasons and landings of razor clams are regulated in Washington. The recreational and commercial seasons are staggered to
prevent illegal sales of recreationally harvested razor clams as commerciallyharvested clams (Ayres & Simons 1992). The recreational bag limit has been reduced from
36–24 to 18–15 clams of any size per digger in Washington (Ayres & Simons 1992). Commercial diggers have no bag limits. In the past, a commercial licence to
harvest razor clams could be obtained for US$5, but many recreational diggers bought commercial licences to avoid the recreational bag limits (Ayres & Simons
1992). Commercial licences now cost US$130 to US$185 and commercial
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diggers must have a Washington State Health Permit, cost US$250. The minimum size for commercial harvest is 114mm shell length and there is no minimum size limit
for recreational fishers (Lassuy & Simons 1989). The autumn and spring harvest seasons are separate and season lengths vary between years and locations (Lassuy &
Simons 1989). Closures of razor clam fisheries due to disease or shellfish toxin accumulation are frequent.
Recreational harvest with minimum size limits produces high associated mortality and wastage of improperly replanted undersized and injured razor clams (Schink et
al. 1983, Lassuy & Simons 1989). In the Washington and Oregon recreational fisheries a regulation requiring that all razor clams must be kept, regardless of size or
condition, has reduced the incidental mortality of undersized clams (Lassuy & Simons 1989). Wastage was high, up to 28% of the harvest in 1949, and has been
reduced to levels of 5–10% (Lassuy & Simons 1989, Ay res & Simons 1992). Despite the new regulations, wastage continues to be a management concern in the
razor clam fishery and has led to the implementation of emergency closures in some areas (Lassuy & Simons 1989). The primary gear used for recreational and
commercial harvest of S. patula is a narrowbladed shovel called a clam gun (Fig. 7); no motorized mechanical harvest is allowed (Lassuy & Simons 1989). Spring
low tides are the best time for intertidal harvest so clamming seasons and enforcement efforts are concentrated around those tides.
In Washington, population assessments of S. patula using markrecapture and stratified random digs are performed at each of the four major recreational clamming
beaches three times per year (Ayres & Simons 1992). Abundance and size structure of populations are assessed before individual beaches are opened to harvest and
these data are used to set the length of the season at each beach. Digger numbers, catch per unit effort, catch per mile of beach and estimates of wastage are made at
each beach during the harvest seasons (Ayres & Simons 1992). The prevalence and intensity of NIX disease in razor clam populations was monitored periodically until
1993 (Ayres & Simons 1992, D.L.Ayres pers. comm.).
Accumulations of phytoplankton toxins in the tissues of razor clams have caused emergency fishery closures and affected fishery landings and economics throughout
the range of the species. Pacific razor clams are tested regularly for the accumulation of toxins, such as PSP and, more recently, domoic acid in many areas. The
commercial razor clam fishery has been closed by high PSP levels in Alaska on occasion (Orth et al. 1975) and high levels of PSP occurred in several razor clam
populations in 1985 and 1987 (Richard Barrett, in Lassuy & Simons 1989). In 1984 high levels of PSP were observed in razor clams in Washington. However, the
fishery was already closed due to NIX disease (Lassuy & Simons 1989). High PSP levels also caused fishery closures on some Washington beaches in fall 1992 and
spring 1993 and led to reduced landings. In 1991 the standard PSP testing revealed domoic acid in razor clam tissue for the first time and the fishery was closed in
Washington (Ayres & Simons 1992, Drum et al. 1993). The Washington razor clam fishery was also closed due to high domoic acid levels for all of 1992, parts of the
fall of 1993 and the spring of 1994 (Ayres & Simons 1994, D.L.Ayres pers. comm.). Periodic closures of the razor clam fishery due to high levels of PSP and domoic
acid have occurred in Oregon. Annual shellfish harvest closures, known as quarantines, occur throughout California to prevent human consumption of toxic shellfish
during the season when toxinproducing phytoplankton blooms may occur.
A number of enhancement efforts have been made on the razor clam resource. Hatchery rearing and release of juvenile razor clams on clamming beaches occurred in
Washington from 1980 to 1989 (Lassuy & Simons 1989). Juvenile razor clams were collected on beaches with high recruitment and transplanted to beaches with poor
settlement in 1966–67 (Tegelberg & Magoon 1969). Rickard & Newman (1986) described collecting juvenile
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Figure 7 a, Collecting Siliqua patula with a “clam gun” in the Pacific Northwest; b, The recreational fishery for S. patula in the
Pacific northwest where fishers number up to 1000 per mile of beach on spring low tides. (Photographs courtesy of the
Aberdeen Daily World).
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clams from the subtidal and transplanting them on to beaches to enhance the fishery. In 1985, 127 million juvenile razor clams were transplanted from the subtidal to the
intertidal zones of beaches (D.L.Ayres pers. comm.).
The Caribbean
Several relatively small tropical species are collected in artisanal fisheries, based on intermediate to reflective beaches in this area.
The guacuco, Tivela mactroides (Fig. 2c, p. 166)
Distribution
T. mactroides is distributed on Caribbean and Atlantic coasts from Mexico to Brazil, including several islands of the West Indies (Fig. 1, p. 164), but it is in Venezuela
where the species has a true commercial value and activity, as a result of the presence of very dense banks of clams in the upper subtidal of finegrained sandy beaches.
Rodriguez (1957) reported such banks for the Maracaibo area (Zulia); Rodriguez (1959), Etchevers (1976) and Prieto (1980b, 1983, 1987), for Margarita Island
(Nueva Esparta) and Eastern Venezuela (Sucre); Brito (1984), Ruiz (1983) and Rincón (1986) for the Barlovento Region in Central Venezuela (Miranda); and
Almeida (1974) for the beaches of Tucacas and Boca de Aroa (Falcon; Golfo Triste area). A number of reports indicate a very high abundance of this clam in the
Golfo Triste area (Almeida 1974, de Mahieu 1984, de Mahieu & Gamba 1980, Penchaszadeh et al. 1979, 1983, Penchaszadeh 1983a, Penchaszadeh & Losada
1987). T. mactroides occurs in dense populations near the mouths of rivers, particularly where large amounts of particles of terrigenous origin are transported in
suspension, or in areas where disturbance of the sediment carries large amounts of material into suspension. Ruiz (1983) reports mean grain size of the sediments from
T. mactroides beaches of 0.145 and 0.195mm (fine sands).
There are probably geographical differences in vertical distribution of T. mactroides related to differences in the tidal regime; on the Atlantic coast of Brazil, T.
mactroides (known locally as “sapinho” or “sapinhagu”) occurs in the intertidal (Narchi 1972) whereas in the Caribbean, the vertical distribution is predominantly
subtidal, to depths of 3m. Here, however the influence of the small tidal range is insignificant when compared with windgenerated changes in nearshore sea level.
Where it cooccurs on Caribbean beaches with Donax denticulatus and D. striatus, Tivela mactroides is found deepest in the sublittoral zone, Donax denticulatus
most superficially and D. striatus intermediate and showing a small overlap with the other two species (Etchevers 1975).

Tivela mactroides is dioecious. Sexual differentiation starts at a shell length of about 12mm and at 20mm length all individuals are sexually mature, with a 1:1 sex ratio
(Brito 1984). Mature gametes are found throughout the year, and there may be many spawning episodes (Prieto 1983). At Playa Guiria, Sucre, Venezuela, the most
intense periods of gonadal maturation were found to be from July–December and April (1974–5) while most spawning activity took place in October–November and
January–February (1982–3) with a minor
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spawning period also in April–May (Prieto 1987). There are monthly variations of the dry weight of the soft parts, mainly associated with reproductive activity; dry
weight for a “standard animal” of 30mm shell length decreases between October–November and January–February, which are important spawning periods (Prieto
1980a). The larval development has not been investigated.
T. mactroides is a suspension feeder, and is often the dominant macrofaunal component of open sand beach communities. The morphological features associated
with this habit have been described by Narchi (1972) and include adaptations of the siphons, ctenidia, labial palps and stomach that enable the clam to exclude sand
particles and to cope with large amounts of material in suspension in the water drawn into the mantle cavity.
T. mactroides are eaten by a variety of predators. Prieto (1987) reported that, at Playa Guiria (Venezuela), juveniles are eaten by the polychaetes Glycera sp.,
Lumbriconereis and Sthenelais boa, and the hermit crab Isocheles wurdemanni, while adults are eaten by the gastropod Natica cayenensis and several fishes
including Cathorops spixi, Caranx hippos, Trachinotus carolinus and Menticirrhus martinicensis. In surveys at Higuerote beach, Central Venezuela in 1987,
remains of Tivela mactroides were found in the stomachs of nine species of fishes; Cathorops spixii, Trachinotus carolinus, Polydactylus virginius, Menticirrhus
littoralis, Ophioscion punctatissimus, Umbrina coroides, Sphoeroides testudinalis, Eucinostomus sp. and Selenopsis sp., reflecting the importance of this bivalve
in the beach food chain (P.E.Penchaszadeh, A.D.Ansell, R.N.Gibson unpubl. obs.). Part of this predation involved nonlethal “grazing” as evidenced by the presence in
the fish stomachs of the severed tips of siphons from the bivalve.
In the Caribbean, living Tivela mactroides shells are used as the substratum for attachment of their egg capsules by Prunum prunum (Gastropoda; Marginellidae)
(Rincón 1986, Penchaszadeh & Rincón 1996). Similarly, on Atlantic beaches the living shells are used as a substratum for the attachment of the eggs of Lintricula
auricularia (Gastropoda; Olividae) (Narchi 1972). Narchi (1972) found large numbers of hydrozoan polyps, probably Eugymnanthea cirrhifera, attached to the
labial palps, ctenidia and visceral mass.
Population dynamics
Populations of Tivela mactroides can reach high densities with a correspondingly high biomass. Prieto (1983) reported mean densities at Playa Guiria, of 788–
1024ind.m−2 with a maximum biomass of 1229gm−2 shell free ww. Brito (1984) found maximum densities of 220ind.m−2 at Higuerote, and 80 ind. m−2 at Paparo. In
Playa La Guardia (Margarita Island) the width of the bank of T. mactroides, as well as the density within the bank, has been found to vary throughout the year
(Etchevers 1976). In general, the bank occupies a sublittoral strip, approximately 70m wide, stretching beyond the beach trough with maximum densities of T.
mactroides in an intermediate fringe located between 20 to 45m from the shoreline. Total biomass (WW of whole animals with shells included) varied from 5.92kgm−2
in November to 12.24kgm−2 in April. Total production for 8.3km of beach was estimated as 3291t, of which some 10% was “noncommercial”, i.e. with shell height
measuring less that 22mm. Prieto (1987) calculated the productivity for Playa Guiria as about 130.07gCm−2yr−1. At Playa Guiria, recruitment of T. mactroides
(<8mm) occurred in November–December 1982 and January, March and June, 1983 (Prieto 1987). This population component also showed rapid growth (3.5mm
month−1) during the first months of life.
T. mactroides has a short life span and rapid growth in those populations that have been studied. The clams grow to a mean shell length of 25mm in 9 months at
Higuerote (Brito
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1984) while at Play a Guiria they reach 30mm in 12 months, with a life span of 18 months (Prieto 1987). Growth may be described by the von Bertalanffy growth
equation: for Higuerote, size frequencies analysis gave values of L∞ and K of 32 mm and 0.23yr−1, respectively (Brito 1984), while for Playa Guiria L∞ was 36mm and
K=0.15 yr−1 (Prieto 1987). values thus ranged between 2.28 and 2.38 (average=2.33: Table 2, p. 168).
Natural mortality is very high; it is common to see many dead and decaying large individuals in the upper surf zone causing the whole beach to smell. As a
consequence of high natural mortality, the majority of the adult population is renewed annually, beginning in July. Using the Van Sickle method (Van Sickle 1977),
Prieto (1987) calculated instantaneous total mortality rates (Z) of 1.56yr−1 (70% annual) for 1974–75 and 2.41yr−1 (91 % annual) for 1982–83. From surveys during
1982–83 of empty shells in which the ligament was still intact (indicating recent mortality), Prieto (1987) estimated an annual mortality of 55%.
Fishery
In common with many other edible molluscs inhabiting shallow waters, T. mactroides has been eaten since preColumbian times. Some cultures also used the shells for
ornaments and trade. Nowadays, a popular dish in Venezuela is “crema de guacuco”, a kind of clam chowder made with T. mactroides meat. Harvesting of T.
mactroides in Venezuela is both recreational and commercial but there are no reliable statistical records for either of these activities. Data collected by the Dirección de
Control de Production of the Servicio Autárquico de Recursos Pesqueros y Acuícolas (SARPA) for the last 10 years show a generally increasing harvest with a
maximum in recorded landings of 3541 in 1992 (Fig. 8). These data almost certainly underestimate the real production, but are the only available source of
information.
Beach clams or chipichipi, Donax denticulatus and D. striatus (Fig. 2d, e, p. 166)
Distribution
D. denticulatus and D. striatus are distributed on Caribbean and western Atlantic shores from Cuba in the north to Brazil in the south (Wade 1967b, Abbott 1974,
Rios 1975, among others) (Fig. 1, p. 164). Both are found in well sorted sand on open sea beaches exposed to wave action, but D. denticulatus occurs on more
beaches and is generally more abundant (Wade 1967b), although D. striatus can dominate and reach high densities in some areas (Etchevers 1975, Farache 1980, De
Mahieu 1984). On the beaches of Barlovento (Venezuela), for example, which are influenced by discharge from the River Tuy, D. striatus is by far the dominant
Donax species, reaching a large size (P.E.Penchaszadeh unpubl. data). D. striatus is a very variable species in colour, shape, and sculpture of the shell and this has led
to some taxonomic confusion in the past (Wade 1967b).
Caribbean beaches have only a small tidal range, seldom exceeding 0.5m, and the vertical distribution of beach clam populations is determined by the physical
conditions at the water’s edge. Etchevers (1975) reported that D. striatus occupies a belt between the upper beach with D. denticulatus and the subtidal Tivela
mactroides population, but in other areas the distributions of the two Donax species may show more overlap, or coincide (Fluck 1906, Wade 1967b, De Mahieu
1984). For D. denticulatus, most of the population is distributed in the wash zone, with only a few individuals in the surf zone, and none in the spray zone.
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Figure 8 Fishery landings of Tivela mactroides and Donax spp. (denticulatus and striatus) from Venezuela (1982–93)
There is also a pronounced sorting by shell size in the wash zone with adults being confined almost entirely to the saturated zone, and only small individuals being found
in the unsaturated zone. These distributions are maintained throughout the tidal cycle and during windgenerated changes in water height by tidal migration (Wade 1964,
1967a,b, Trueman 1971, Ansell & Trueman 1973).

Both D. denticulatus and D. striatus are dioecious. For D. denticulatus, Velez (1985), studying a population on Araya beach, Venezuela, reported that
approximately 3% of juveniles of shell length 7–9 mm showed signs of gametogenesis although it was was difficult to determine the sex of individuals smaller than
11mm. The number of sexually mature individuals increased progressively to the 17–19mm group, in which 98% could be sexed. Juveniles and adults showed a 1:1 sex
ratio. In this Venezuelan population, reproduction continued throughout the year although with some seasonal variation in the proportion of mature and spawning stages
in the population. There was greater reproductive activity during July– December. For the populations of D. denticulatus he studied in Jamaica, Wade (1968)
concluded that reproduction was seasonal, with ripe gonads being found mainly between August and January.
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Wade (1968) found veliconcha stages of D. denticulatus in the plankton throughout the year in Jamaica, but they were especially abundant from November to
January. Settled spat were found in every month over a period of nearly 2yr, but, nonetheless, a very well defined peak of settlement could be observed beginning in
November. Similarly, Vélez (1985) reported monthly recruitment of D. denticulatus (of spat measuring 0.1 to 4.0mm) with minimum densities from January to June,
and maximum densities from July to December. During this period there were peaks in September, October and December 1974, coinciding with observed gonadal
changes. Wade (1968) reported that veliconchas of D. denticulatus taken from the plankton and reared in the laboratory lived for up to 3wk during which they
sometimes doubled their size. He figured a late veliconcha of D. denticulatus but did not describe the larval development in detail.
Both D. denticulatus and D. striatus are important in the beach food chain as abundant primary consumers of phytoplankton and detritus. In turn these clams are
eaten by a wide range of predators throughout the range of distribution. In the West Indies, Wade (1967a) lists portunid (Araneus cribarius and Callinectes
exasperatus) and ocypodid (Ocypode albicans) crabs, and birds (the sanderling Crocethia alba), as predators on Donax denticulatus together with the pig: “The
pig’s appetite for Donax is immense, and it seems that one pig in a day may devour more than what all other predators consume in one month”. In Venezuela, D.
denticulatus is eaten by a variety of fishes including Menticirrhus littoralis, Conodon nobilis, Trachinotus carolinus, T. goodei and Umbrina coroides
(Penchaszadeh 1983b, Riera 1995, A.D.Ansell, R.N.Gibson & P.E.Penchaszadeh, unpubl. data), and by ghost crabs Ocypode quadrata and the gastropods Prunum
prunum and Olivella verreauxi. Menticirrhus littoralis and Trachinotus carolinus were found to have eaten both entire Donax and in some cases only the siphons.
Wade (1967a) recorded a low level of infestation of a bucephalid cercarian trematode Cercaria caribbea XLII in Donax denticulatus from Port Henderson and
Green Bay, Jamaica. Parasitic castration had occurred in all the affected clams that appeared otherwise healthy. The shells of D. denticulatus are also used as a
substratum for attachment by hydroids and algae (Wade 1967a).
Population dynamics
Population densities of D. denticulatus and D. striatus are variable both between locations and at different times at one site. For D. denticulatus, Wade (1967a)
recorded densities ranging from 10 to 4120ind.m−1 of beach on 22 beaches in the West Indies. At one site, Farquhar’s Beach, Jamaica, the population density fell from
4120 to 360ind.m−1 over an 18 month period. Recorded maximum population densities for D. denticulatus on beaches in Venezuela range up to 2150ind.m−2 for
populations consisting mainly of juveniles and up to 509ind.m−2 for adults (Farache 1980, Penchaszadeh et al. 1983, Diaz et al. 1987, Pauls et al. 1990). Etchevers
(1975) evaluated the biomass of D. denticulatus in Playa La Restinga, Margarita Island, Venezuela and estimated 14.2t (ww including shells) in 18km of beach.
For D. striatus, Wade (1967b) reported a density of 3000ind. m−1 for a beach in Trinidad. In Venezuela, Farache (1980) reported a mean density of between 3.6
and 208ind.m−2 with a maximum of 496 ind. m−2 and De Mahieu (1984) found a maximum number of 1264ind.m−2 at Tucacas beach, Venezuela, where Pauls et al.
(1990) reported a maximum adult density of 198 ind.m−2, and a juvenile density of 509ind.m−2.
Age and growth for D. denticulatus were estimated by Velez et al. (1985) from size frequency distributions of clams collected from a beach of the Araya Peninsula,
Venezuela.
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A similar recruitment pattern occurred in both the years studied, 1974–75. Recruits, i.e. juveniles less than 4mm total length, appeared continuously through the year,
but were most abundant between July and December. A distinct juvenile cohort (modal length 8–9mm) could be identified in June 1974 and remained clearly
identifiable in the subsequent histograms for that year. August–September recruits first became noticeable in the juvenile size classes of the October–December 1974
histograms. By January 1975, the cohort of September 1974 recruits had become the dominant component in the population. Its decline by November 1975 to
February 1976 suggested a life expectancy of only 13–16 months. Individual cohorts were less distinct in 1975–76 than in 1974–75 but in general, there was a
dominance of juveniles from January through July and a dominance of adults from August through December. The rate of increase in total length was relatively high (4.2
mm month−1) during the first 2–3 month post recruitment. It fell to 1mm per month during the next 4 months and after 7 months appeared to be negligible. The mean
maximum length did not exceed 19mm in this locality.
The von Bertalanffy growth equation calculated for this population of D. denticulatus (Vélez et al. 1985) was:
where Lt is shell length in mm at time t in months and t 0 value (0.058) is the theoretical age when the mean clam had zero length. The maximum size reached by D.
denticulatus in different populations varies greatly depending on conditions on different beaches. Wade (1967a) reported maximum sizes ranging from 12–23 mm for
different populations in Jamaica.
Population densities for the population studied by Velez et al. (1985) ranged over approximately one order of magnitude during the 2 years of study, from 212 to
2435ind.m−2. Population density of individuals 4mm or less in shell length increased markedly from March to July principally from growth of recruits during November
to December while mortality resulted in decrease in density of individuals of 4mm or more in shell length from August to November.
Estimated daily production ranged from 31 to 466 mgm−2, with standing dry biomass values of 4 to 40 gm−2. Integration of daily production over both years gave
annual production estimates of 37.6gm−2 and 77.4gm−2 for 1974–75 and 1975–76 (equivalent to 181 and 372kcalm−2).
For D. striatus, Farache (1980), found two recruitments, one in March–April, and another in August, during a 9month period of sampling at Tucacas and
surrounding beaches in Venezuela. The growth rate at these beaches was very rapid during the first months after the recruitment, the clams reaching 50% of their
asymptotic size in 2½ months, 66% in 4 months and 90% in 8 months. The mean monthly growth rate was 2.04mm. The von Bertalanffy growth curve for D. striatus
at Tucacas (where La=the distance from the umbo to the end of the anterior part of the shell) was:
As with D. denticulatus, the maximum size reached by D. striatus varies greatly between areas and at different beaches in the same area (Wade 1967b). The
maximum size of clams in the population studied by Farache (1980) was relatively small.
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Fishery
The official statistics of landings of Donax in Venezuela probably greatly underestimate the catches and do not discriminate between D. denticulatus and D. striatus.
Adult individuals of D. striatus generally reach a larger size, however, and consequently this is the preferred marketable species. Data on landings (expressed as kg
weight including shell) for the 12 years 1982–93 are given in Figure 8 (p. 183). Recorded landings have been variable with a maximum of 226t in 1990.
Additional studies of D. denticulatus and D. striatus are necessary to provide information on geographic and amongyear variation in density, growth and
production, and the effect of exploitation. At this time insufficient data are available for longterm fishery management decisions. Specifically, stock recruitment
assessments would help to determine the susceptibility of the Donax populations to overexploitation. Based on the results of Vélez et al. (1985) and Vélez (1985),
however, there is the possibility that population stability could be sensitive to fishery exploitation of commercialsize individuals during the peak period of reproduction.
Thus, Vélez et al. (1985) recommended that fishing should be limited to the period November–April in Venezuela.
Southern South America

A number of sandybeach bivalves are exploited through recreational, artisanal and commercial fisheries on both the Atlantic and Pacific coasts of South America. Only
the two most important species are covered here.
The yellow clam, Mesodesma mactroides (Fig. 2f, p. 166)
Distribution
M. mactroides is an intertidal bivalve distributed along the warmtemperate Atlantic coast of South America, from Sao Paulo State, Brazil (24°S) to the south of the
Buenos Aires Province, Argentina (41°S; Fig. 1, p. 164). This range includes hundreds of kilometres of sandy beaches in Brazil, 22km in Uruguay and beaches of the
Buenos Aires Province in Argentina (Olivier & Penchaszadeh 1968a). This review is based mainly on the yellow clam population in Uruguay, which has been the most
extensively studied.
Yellow clam populations prosper primarily in the intertidal zone of dissipative beaches (sensu Short & Wright 1983) characterized by gentle slope, fine sand and
heavy wave action (Olivier et al. 1971, Defeo 1985a, Defeo et al. 1986). A decrease in clam abundance is frequently observed near estuarine zones or freshwater
discharges (Defeo et al. 1986) and on sheltered beaches (Defeo et al. 1992a). The beach that M. mactroides inhabits in Uruguay is microtidal dissipative (slopes
3/100, fine to very fine sands) with high wave energy and a wide surf zone. It is characterized by high primary production by surf diatoms (Defeo & Scarabino 1990)
that allows the beach and surf zone to function as a semiclosed ecosystem (sensu McLachlan 1980).
M. mactroides presents a marked longshore distribution in the 22km beach; abundance is highest in the central zone and declines towards both ends. Freshwater
inflows, coarser and poorly sorted sands (negatively skewed and high kurtosis values), the dominance of erosional
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processes, and a selective winnowing of fine sediments, contribute to a reduction of the stock towards the ends of the beach (Defeo 1993a). Salinity is positively
correlated with clam abundance, largely explaining the alongshore distributional pattern.
The intertidal and vertical distribution of M. mactroides varies seasonally (Coscarón 1959, Olivier et al. 1971, Defeo et al. 1986). In winter, adults (ages 1–4) are
found 30–40cm deep in the sediments and below the swash zone, while in summer they are located close to the sand surface and almost exclusively above the swash
zone. In spring and autumn, intermediate situations are commonly observed. Water temperature and correlated factors (e.g. chlorophyll concentration, gonadal
development) appear to be important in determining the intertidal distribution and depth stratification of M. mactroides adults. Recruits are mainly concentrated at
upper intertidal levels (Defeo 1985a, Defeo et al. 1986). These spatial gradients for recruits and adults suggest a differential capability of each population component to
select the desirable microhabitat. Whereas recruits appear to be more dependent on passive transport (waves, tides), seasonal variability in the zonation of adults may
be the result of an active process which involves a certain level of mobility and a complex set of behavioural adaptations (e.g. burial depth with an anchoring system; see
Narchi 1981) to variations in environmental factors such as temperature, sand desiccation and food availability (Olivier et al. 1971, Defeo 1993a).
Temporal variations in the mean clam bed width (CBW) can be explained for adults and recruits by an asymptotic function of density (Defeo 1993a). Asymptotic
values of the mean CBW were close to 13m. Oliver et al. (1971) also mention a clam bed width ranging from 4m to greater than 20m, depending on the beach face
slope. Distribution is aggregated and patch characteristics show a clear trend toward ellipses, with the major axis parallel to the shore (Defeo et al. 1986, Defeo
1993a). Within each patch, a concentric gradient may arise from (a) passive transport and sorting by the swash (e.g. recruits), (b) similar reactions to environmental
factors, and (c) active selection of a suitable microhabitat to avoid predators or to search for localized food concentrations (e.g. adults).
Distinct aggregation of all ages occurs throughout the year. Results obtained from the analysis of the variance: mean ratio (even within a year) showed the capacity of
M. mactroides to develop high densities. The patchy distribution among recruits and the inverse relationship between individual size and density (Defeo 1993a),
suggest that the clams tend to space themselves as a function of an increasing area of influence needed for filtering activities (see also Hall 1983).

A 1:1 sex ratio and two spawning periods were observed for Uruguayan (Defeo 1985b, Masello 1987) and Argentinian populations (Olivier et al. 1971): one between
October and December and a second one from February to April (late summer). Sexually ripe individuals of 2–4 yr are responsible for the former, while newly mature
individuals of age 1 (from 40mm to 45mm) spawn in late summer (Masello 1987). Gonadal development is controlled by temperature, with spawning taking place
when the temperature is close to 20°C. M. mactroides has a high fecundity, producing about 5.3 million eggs per female. The mean size at first maturity is 43 mm (age
1; Masello & Defeo 1986). Settlement occurs in the intertidal zone from January to May.
M. mactroides has been commonly described as a suspension feeder (Olivier et al. 1971, Narchi 1981, Gianuca 1983, Defeo 1985a). Diatoms such as
Asterionella glacialis, the basic food source (Gianuca 1983; Defeo 1985a), Skeletonema costatum, Thalassionema nitzschoides and Coscinodiscus spp.
constitute the bulk of the stomach contents. Laboratory and field observations (Defeo & Scarabino 1990) showed that it also has a depositfeeding mechanism.
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Some special features of the adaptive morphology of the species (Narchi 1981), attributed to an adaptive divergence of the superfamily Mactracea (Stanley 1970),
could support the depositfeeding habit (Defeo & Scarabino 1990).
Mesodesma mactroides is the dominant suspension feeder in the intertidal macrofauna assemblages of dissipative sandy beaches in Uruguay (Defeo et al. 1992a),
Brazil (Gianuca 1983) and Argentina (Olivier et al. 1971). Another potentially exploitable bivalve, the wedge clam Donax hanleyanus (Philippi), is sympatric and
subdominant to Mesodesma mactroides in those beaches (see also de Alava 1993). Donax hanleyanus is numerically dominant on the reflective beaches with coarse
sediments and sharp slopes along the Atlantic coast of South America, from the state of Espiritu Santo (Brazil) (Narchi 1978) to Punta Mogotes, Buenos Aires
Province (Argentina) (Penchaszadeh & Olivier 1975). This intertidal filter feeder has a maximum adult size of 35mm, and rarely lives for more than 3yr (Penchaszadeh
& Olivier 1975).
Competition between Mesodesma mactroides and Donax hanleyanus has been indicated by a longterm study in Uruguay (de Alava 1993, Defeo & de Alava
1995). Spatiotemporal variations in recruitment success and population levels of D. hanleyanus were inversely correlated with the magnitude of fishing effort exerted
on Mesodesma mactroides. This human impact caused a reduction of M. mactroides stock and a consequent release of space that enhanced recruitment of Donax
hanleyanus. Thus, even though Mesodesma mactroides is a suspension feeder, it may be a keystone species in the sense that fluctuations in its magnitude can generate
cascade effects affecting the structure and abundance of the sympatric species.
The main predators of M. mactroides are the gastropods Olivancillaria vesica auricularia and Buccinanops duartei, fishes like Micropogonias furnieri,
Paralichthys brasiliensis, Mustelus schmiti and others of the genera Raja and Psammobatis, and birds such as the oystercatcher Haematopus ostralegus and
Rinchops nigra (Olivier et al. 1971). The large cirolanid isopod Excirolana armata has been observed feeding on damaged, stranded or unburied clams.
Population dynamics
The yellow clam population in Uruguay displays temporal “stockdependent recruitment” (sensu Hilborn & Walters 1992). A significant overcompensatory
relationship between spawning stock and recruitment of Mesodesma mactroides was modelled for the first time in an exposed sandy beach population by Defeo
(1993a). Two definite trends emerged: (a) larger adult densities produced extremely low recruitment and (b) maximum recruitment densities occurred from moderately
low and medium sizes of spawning stock. The proposed mechanism of presettlement, densitydependent mortality is the filtering of larvae out of the water column by
the adult stock.
The relation between stock and recruitment varies with the scale: even though overcompensation exists at a macroscale, transect and site analyses show a consistent
pattern of alongshore distribution common to adults and recruits: the central portion of the habitat is more densely populated than the margins, whereas zones with low
adult density receive fewer recruits. Higher mortalities before and after settlement because of adverse environmental effects (i.e. lower salinities), could account for this
variability. At a quadrat scale, high densities of recruits never coincided with high densities of older clams. Recruitment success appears to be highly inhibited in
quadrats with high density and biomass of adults. The recognition of a certain maximum width of the clam bed suggests a limitation of avail
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able space at an adult density close to 120m−2. This value is consistent with those suggested by the spatiotemporal analysis of the stockrecruitment relationship
(Defeo 1993a).
The yellow clam is a fastgrowing shortlived species (Table 2, p. 168). Maximum observed length ranged from 78mm (Uruguay) to 80–85 mm
(Argentina: Carcelles 1939, Olivier et al. 1971). Growth parameters have been estimated by methods that use age or length data in Uruguay. Age structure, determined
by the interpretation of growth rings and by shell sectioning techniques, suggests a life span of ≈3.5yr (Table 2). Rings of different periodicity are identified: “winter
rings”, formed during late fall and winter, coinciding with the period of null growth determined by lengthfrequency analysis; and secondary rings during the main period
of gamete production, considered as “reproductive rings” (Defeo et al. 1988a). Growth rates are higher during the summer season, dropping between fall and winter,
and increasing from September (austral spring) (Defeo et al. 1992b). A 14°C difference between minimum and maximum mean monthly values of temperature in the
study area (de Alava & Defeo 1991) accounts for the seasonal variation in growth rates. Densitydependent growth rates were demonstrated in a longterm study
(Defeo 1993a).
Olivier et al. (1971) did not report seasonal variations in growth for clams in Argentina, but estimated lower growth rates (i.e. K≈0.25–0.29yr−1) and greater
longevity (≈8yr) than for studies in Uruguay (Defeo 1993a). The lack of seasonality was probably due to the growth model employed (von Bertalanffy 1938), whereas
lower growth rates could be attributed to the assumption of annual periodicity in the formation of rings (without this being corroborated by any other method).
However, differences in growth rates may be a response to dissimilar environmental characteristics. Defeo (1993a) reported lower sizes at marginal, less favourable
sites, with a virtual absence of clams of 40mm length. Considering these effects over 22km, growth rate differences within the geographical range of the species are
likely.
Estimates of the instantaneous rate of total mortality (Z) were obtained by means of catch curves drawn according to the agelength key, and had values of 1.81yr−1
and 2.55yr−1 for 1983 and 1984, respectively. From age 2 on, the clams were fully available to the catch (Defeo et al. 1992b). The increase in Z for 1984 could be
attributed to an increase in densitydependent natural mortality (i.e. greater densities for the 1984 cohort), or fishing pressure or both. However, a high variability in
mortality rates among years implies that the above values must be treated with caution (see below).
The instantaneous rate of natural mortality (M) varied. For the closed fishing season 1988–90 an M value of 1.64yr−1 was obtained from changes in abundance of
the cohort over time. Based on the unexploited fraction of the population (1–50mm) for years 1983 to 1987 and 1990, when the fishery was open, M ranged from
0.90yr−1 (1990) to 3.17yr−1 (1985). M was highest in the densest cohorts, which also exhibited lowest growth rates under intense fishing pressure (Defeo 1993a). M
also varied spatially, according to the quality of the site, the extent of adultrecruit interactions and the heterogeneous allocation of fishing effort. A longterm study
between 1983 and 1990 detected potential incidental mortality in the youngoftheyear, produced by damage and sediment disturbance during harvesting activities: M
was significant and positively correlated with the amount of fishing effort and catches observed per fishing ground (Defeo 1993b). Fishing mortality values ranging from
0.62yr−1 for 1983 to 1.27yr−1 for 1984 were obtained by subtracting M from Z. A cohort analysis was useful to determine the extent of fishing mortality per age group
(Defeo et al. 1988b).
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Fishery
Information on the M. mactroides fishery in Brazil is scarce, and it can be considered an openaccess system. The yellow clam can almost disappear at the end of each
summer season around every important seaside resort, where it is consumed as food and utilized as bait (Gianuca 1983). In Argentina, exploitation of M. mactroides
has been carried out since the 1940s, concurrently with the development of the canning industry (Olivier et al. 1971). Landings peaked in 1953 at 1078t, then declined;
a total closure of the fishery has been implemented since 1958 (Olivier & Penchaszadeh 1968b). To date, even with the fishery closed, the stock has diminished due to
illegal extraction, including that associated with summer tourist activity (Bastida et al. 1991).
In Uruguay, the yellow clam constitutes an important artisanal fishery. The clams are collected by handgathering techniques and marketed either for bait or human
consumption, mainly during the summer months and in the seaside resorts. Prices paid to fishers range US$0.5–1kg−1, and market value for consumers is near
US$5kg−1 (including valves). The enhanced demand for the clams caused a fast growth in landings during the 1980s, ranging from 621 in 1981 to a peak of 219t in
1985 (Fig. 9). Thereafter landings decreased rapidly. A temporal closure of the fishery was imposed between April 1987 and December 1989.
An overall catch quota, the first management measure implemented, was unsuccessful; rapid collection and stockpiling of organisms meant that the catch quota was
quickly reached, leading to a deterioration in the product. A minimum profitable catch volume per fisher, differential fishing quotas per season and rotation of areas
were therefore recommended as tools for the management of the fishery (Defeo 1989). For the implementation of such strategies, habitat heterogeneity, which accounts
for spatial and temporal variations in resource abundance and in the fishing effort exerted, was considered. A minimum legal size of 50mm (≈1.5yr) for commercial
harvest, based on the length at first maturity, is also used as a regulatory measure (Defeo 1987).
Management of the Uruguayan stock was improved through experimental procedures that included natural repopulation of depleted areas, rotation of areas and the
allocation of indi
Figure 9 Time series of Mesodesma mactroides landings, 1981–91. (After Defeo et al. 1993).
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vidual quotas per fisher through a catch per month per fisher. This number was determined as the ratio between the net value of the overall quota and twice the annual
minimum salary (Defeo 1993b). To implement the spatial management plan, fishing grounds with lower productivity were used as exploitation units when demand for
clams was low (i.e. austral autumn and winter), whereas the central sections, with highest stock and recruitment densities, were used in spring and summer (Defeo et al.
1991). This period maximizes the biomass of the cohort available for harvesting (Defeo et al. 1992b). Rotation of fishing areas also maintains space and a certain level
of adult density to enable recruitment success at desirable levels.
The wide range of levels of fishing effort achieved by experimental management provided a range of variability in the spawning stock size, and allowed determination
of population patterns and processes that were obscure before closure, for example the exact shape of the recruitment curve. The spatially disaggregated analysis of the
resource and the fishery constituted a useful approach in interpreting the behaviour of this exposed sandybeach population (Defeo 1993a). Results of the longterm
study in Uruguay demonstrated that quantitative models developed for exploited populations from sandy beaches must take into account densitydependent, or area
specific variations in growth, recruitment and fishing and natural mortality (Defeo et al. 1993).
Simulation of alternative management strategies (Defeo 1993a) suggested that, if the population is closed to fishing activities for a long period, high adult density
could inhibit recruitment success and hence the magnitude of the stock available for fishing. However, a closed season of 2 years could promote successful recruitment
in the short term, and an increase in population growth above that observed under an exploited scenario. Fishing pressure acts as a source of disturbance, but also as a
way of releasing space, thus enabling the recolonization of disturbed patches. Harvesting activities within certain levels may diminish the magnitude of presettlement
natural mortality and promote successful recruitment. Because of very low levels of the unit cost of effort, bioeconomic equilibrium occurs at high levels of fishing
intensity, which in turn increases the risks of exhausting this highly vulnerable sedentary species.
The macha, Mesodesma donacium (Fig. 2g, p. 166)
Distribution
M. donacium is distributed along the Pacific coast of South America, from Bahía Sechura (Perú, c. 5°S) to Isla de Chiloè (c. 43°S) (Tarifeño 1980) (Fig. 1, p. 164).
This species inhabits the swash zone to about 15–20m deep off exposed highenergy intermediate and dissipative sandy beaches (Tarifeño 1980, 1984). It has also
been found in sheltered sites in the outlets of estuarine areas of south central Chile (E.Jaramillo unpubl. data). The acrossshore distribution of M. donacium is patchy
and adult clams are primarily confined to the surf zone, while the vast majority of juveniles occur in the swash zone (Tarifeño 1980, Jaramillo et al. 1994).
The longshore distribution of M. donacium is highly variable (Tarifeño 1980, Jaramillo et al. 1994). For example, in a sandy beach stretching approximately 1800m,
several surf clam beds were clearly distinguished; patches of surf clams are usually separated by areas with very low densities or by vacant areas whose lengths can
reach several hundreds of metres (Jaramillo et al. 1994). The discontinuous distribution of this species might be related to longshore differences in grain size (Tarifeño
1980) and/or variability in compactness of the substratum (E.Jaramillo unpubl. data).
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Populations of M. donacium usually have a sex ratio of 1:1 (Salgado & Ishiyama 1979, Tarifeño 1980, Peredo et al. 1987). Studies of the reproductive cycle of M.
donacium in Peruvian and Chilean waters have shown differences in the timing of gametogenesis and spawning periods along this latitudinal gradient. These differences
may be related to local variability in environmental factors such as water temperature and food availability (Tarifeño 1980, Peredo et al. 1987). Surf clams matured in
Peru (≈16°S) between late winter and late spring (August–November), while the spawning period lasted from late spring to midwinter (December–July), the last stage
being most frequent during summer months (December– March) (more than 85%) (Salgado & Ishiyama 1979). In latitudes close to 33°S, ripe animals peaked (95%)
in midwinter (August), while spawned surf clams were more frequent in September, with less than 20% in this stage until late summer (March) (Tarifeño 1980). Surf
clams collected further south (≈37–38°S) were mature during winter, while the spawning season was primarily during late spring (November–December) (Tarifeño
1984); Tarifeño also mentioned a second spawning period (late summer; February–March). Clams of south central Chile (≈39°S) had a maturation period during
winter and spring (from June through November), ripe animals being most abundant in November (Peredo et al. 1987) and spawning extended from summer to early
autumn (December to April). Further south (≈40°S), M. donacium has two main periods of spawning: one during spring (October–November), and the second one
during late summer (López 1976). The duration of larval stages is unknown, as are the dynamics of recently settled larvae and recruits. However, in central and south
central Chile, recruitment takes place primarily at sheltered sites near estuarine outlets (Tarifeño 1980; E.Jaramillo unpubl. results).
Size at first maturity (Table 2, p. 168) also varies geographically in M. donacium, but there is no clear latitudinal pattern. Maturation occurred at 35–70mm shell
length (mean 56mm) in Peru (≈16°S) (Salgado & Ishiyama 1979), and at 16–18mm in central Chile (≈33°S) (Tarifeño 1980). Further south, maturity occurred at 28–
76mm (mean 49.1mm) (Tarifeño 1984), while López (1976) mentions a range of 47–57mm.
Population dynamics
Age and growth of M. donacium have been studied by analyses of lengthfrequency histograms and growth rings on the shell (Alarcon 1979, Tarifeño 1980, 1984,
Arntz et al. 1987). Comparisons of results showed that growth rates vary with latitude (Table 2, p. 168). In Perú (12°S), Arntz et al. (1987) calculated a maximum
shell length of 110–124mm and annual growth rates value for the species throughout its entire distribution range was close to 3.74.
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Fishery
Landings of M. donacium in Peru showed a steady increase from 1964 (36t) to 1977 (597t), reaching an impressive peak between 1978 and 1979, when catches
were sevenfold higher (≈4000t: see Fig. 10). This happened during the period preceding the ENSO event of 1982–83. The decline from 1980 to 1985 may be a
combined result of resource overexploitation (1980–81) and negative effects of ENSO on M. donacium populations (Castilla & Camus 1992). Arntz et al. (1987)
pointed out that from March 1983 onwards, no live surf clams were found in shallow waters south of Lima and reported no recovery up to 1986. Landings from 1986
to 1989 showed a recovery of the resource, but never reached the landing levels of 1978–79 (see Defeo et al. 1993).
Figure 10 Mesodesma donacium landings in Chile (1955–91) and Peru (1964–89). (After Defeo et al. 1993).
In Chile, surf clams are primarily harvested by artisanal fishers operating from the beach and entering the surf zone during low tides. However, most of the catch is
ultimately sold on commercial markets. The fishers operating from the beach, called “macheros”, collect the clams by twisting their feet and using their body weight to
excavate the sand until clams emerge at the sediment surface to be picked up by hand. In some areas (especially during very calm days) M. donacium is also harvested
by hookah divers from boats located behind the surf zone.
The exploitation of M. donacium is regulated only by a minimum size of 50 mm shell length in south central Chile (≈38–42°S) and 60 mm shell length in the rest of
the country. Landings data gathered from Government sources (Servicio Nacional de Pesca, SERNAP) between 1965 and 1991 show a marked increase from 1983
onwards (Fig. 10), as a response to a strong trend towards diversification in the export of Chilean shellfish products (Defeo et
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al. 1993). Landings peaked in 1989, reaching c. 18000t, and declined thereafter (1990–91) down to 9000t (Fig. 10), probably due to overexploitation and/or
fluctuations in market demand. Indeed, a trend of the foreign market (i.e. Spanish market) towards the selection of clam sizes lower than the legal marketable size, has
been observed (Defeo et al. 1993). According to these fishery statistics, areas located in south central Chile account for the majority of the landings. However, not all
the catch is reported to SERNAP; often the macheros use the catch for their own consumption, or else sell it directly to consumers.
A daily set of data on the fishery of M. donacium was collected from May 1992 to October 1994, on a 2km stretch of beach in south central Chile (Mehuín, c.
39°S). During this period, the maximum numbers of macheros working in a single day were 73, 23, and 20 (February, April 1993, and March 1994, respectively). The
average maximum number of fishers was three macheros per day. Open harvesting seasons for other species (primarily “loco” Concholepas concholepas) influenced
the number of fishers collecting surf clams during some stages of this study. The maximum number of working days (up to 90% of the whole month) occurred during the
austral summer months of January–March, whereas during winter and early spring months few days were worked (less than 40% of the whole month), primarily
because of rough surfzone conditions. The total catch ranged from 58–6558kg per month, and peaked during summer months. Lowest harvests occurred during
winter months, for example less than 100kg per month during JulyAugust 1994. However, the mean catch of surf clams per machero remained fairly stable, (i.e. 10.5
kgday−1).
Europe
Only one species of clam is widely harvested on exposed sandy beaches in Europe, although many are utilized on tidal flats and in estuaries.
The wedge clam, Donax trunculus (Fig. 2h, p. 166)
Distribution
D. trunculus is distributed along the Atlantic coasts of Europe and North Africa, from the Bay of Douarnenez, Brittany, France (Guillou & Le Moal 1980) to Senegal
(Fischer et al. 1981), and throughout the Mediterranean and Black Seas (Fig. 1, p. 164). Within this range of distribution, abundant populations are found in areas of
high productivity often, but not exclusively, associated with the discharge of rivers. Distribution is limited to moderatelyexposed or exposed beaches of clean, well
sorted sand with median particle sizes in the range 190–370μm (Mazé & Laborda 1988, Guillou & Bayed 1991). For beaches along the coast of Morocco, there is a
close correlation between the maximum density of adults found and the distance from the coast to the 30m depth contour, indicating dependence of density on
hydrodynamics which is controlled by the gradient of the sea bed inshore of this critical depth at which wave energy begins to be dissipated (Guillou & Bayed 1991).
Hydrodynamic conditions also account for differences in population density along the length of beaches (Mouëza & Chessel 1976).
The vertical distribution of D. trunculus varies in different parts of its range. In the essentially tideless Mediterranean Sea, D. trunculus occupies a relatively narrow
zone in depths of
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a few centimetres to 1m (Massé 1971, 1972, Amouroux 1972, Degiovanni & Mouëza 1972, Geldiay & Uysal 1972, Mouëza 1972a, Bodoy & Massé 1979, Salas
Casanova 1987) with a second smaller species of Donax, D. semistriatus, occupying a similarly narrow zone in slightly deeper water (Massé 1971, 1972) and a third
species, D. venustus, yet deeper. On the tidal Atlantic coasts, D. trunculus occurs in the intertidal zone (Salvat 1968, Chassé 1972, Gruet 1974, Guillou & Le Moal
1978, 1980, Ansell & Lagardère 1980, Guillou 1982, Otero 1982, Bayed & Guillou 1985, Guillou & Bayed 1991) usually with the size classes partially segregated;
juveniles occupy the higher levels while increasingly larger individuals are found closer to low water; the largest individuals occur near low water mark of spring tides.
In the northern parts of its range on Atlantic coasts, the distribution of D. trunculus overlaps the southern part of the range of a second smaller species, D. vittatus.
Here the vertical distributions of the two species also overlap, but the latter reaches its greatest population density subtidally (Guillou & Le Moal 1978, 1980, Ansell &
Lagardère 1980, Guillou 1982). Near the southern part of the range of D. trunculus, D. venustus occupies a similar essentially subtidal zone (Guillou & Bayed 1991).
In the Bay of Malaga, Spain, just within the Mediterranean Sea, four Donax species are all present in the same area with overlapping vertical distributions; here D.
trunculus are the most superficial, D. vittatus and D. semistriatus slightly deeper, and D. venustus the deepest (Salas Casanova 1987).

In D. trunculus the sexes are separate and the sex ratio does not differ significantly from 1:1 (Lucas 1965, C.Tirado & C.Salas, pers. comm.). There is no sexual
dimorphism of the shell, but males and females can be readily distinguished on dissection by the colour (white and green, respectively) of the gonad, except during the
inactive period of the annual reproductive cycle.
The reproductive cycle of D. trunculus shows essentially the same characteristics throughout its range of distribution (Lucas 1965, Mouëza & Renault 1971, Badino
& Marchionni 1972, Poli 1972, Mouëza & FrenkielRenault 1973, Ansell & Bodoy 1979, Ansell et al. 1980b, Bayed 1990). Following an “inactive period” or
“resting stage” of variable duration depending on the geographical area, gametogenesis begins either in the autumn or winter. Spawning first takes place in the spring
and this is followed by further spawning(s) before the reproductive cycle is completed in late summer or autumn and animals have returned to the resting stage. During
the resting period, the gonad is regressed and the sexes are indistinguishable macroscopically. There may be two major spawnings during the year; in June/July and
September/early October on the Camargue coast of France (Ansell & Bodoy 1979) and in spring and summer on the Moroccan coast (Bayed 1990), or spawning
may be more evenly distributed throughout the summer, as on the Algerian coast (Ansell et al. 1980b). On the coast of Málaga, southern Spain, spawning was found to
be continuous from April to October but with peaks in May and August (C.Tirado & C Salas pers. comm.). The start and duration of the different stages shows some
regional variation related to differences in the annual cycle of seawater temperature (Bayed 1990), or with the cycle of phytoplankton production (C.Tirado & C.Salas
pers. comm.).
The tissue weight and the biochemical composition of the tissues show characteristic seasonal changes associated with the reproductive cycle and with the associated
cycle of storage and utilization of reserves in the tissues (Ando et al. 1976, Ansell & Bodoy 1979, Ansell et al. 1980b, Dowidar & ElNady 1984). On the coast of
Algeria (Ansell et al. 1980b), the dry
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tissue content is low in winter and rises rapidly in the spring and early summer as the gonad increases in size. Dry tissue content then fluctuates depending on the balance
between gametogenesis, storage of reserves and spawning, but falls again at the end of the reproductive period. The nitrogen (protein) and lipid content reflect these
changes. On the Camargue coast of France (Ansell & Bodoy 1979), the cycle is similar to this in many respects but there is a rise in tissue content in late autumn,
following the fall that marks the end of the reproductive period, probably reflecting a higher food (phytoplankton) availability at this time in this area.
For D. trunculus from the coast of Málaga, Southern Spain, where there are two peaks of spawning, C.Tirado & C Salas (pers. comm.) estimated that fecundity
(total number of oocytes contained in the ripe gonad) ranged between 58900 and 641000 for females collected in May and between 33245 and 361000 for females
collected in August and September. At the time of spawning, the gametes are released into the water where the eggs are fertilized. Development has not been described
in detail for D. trunculus but Otero (1982) reared larvae through to settlement in the laboratory and his results indicate that development is similar to that described in
more detail for D. vittatus (Frenkiel & Mouëza 1979). The spherical eggs (70–80 μm diameter after fertilization) develop through freeswimming blastula and
trochophore stages to produce a Dlarva (86–95 μm length×78–85 µm height in D. vittatus) after 48h. Mouëza (1975) figured a fertilized egg of 76μm diameter and a
Dlarva of 100 μm length×80 µm height for D. trunculus, but did not succeed in rearing larvae to settlement. Otero (1982) figured a Dlarva of 110×90 μm. Further
development is typically planktotrophic. Growth of the veliger larva is presumably dependent on sea water temperature and the availability of suitable microalgae as
food as in other bivalves. Settlement and metamorphosis takes place at the pediveliger stage that, in D. vittatus, is reached at a shell length of 250–350 |iim after some
31 days at a temperature of 16–18°C. For D. trunculus, metamorphosis took place after 19–23 days at 21°C (Otero 1982). Neither settlement behaviour, nor post
settlement redistribution of juveniles has been studied in D. trunculus, but there is a strong supposition from the patterns of distribution of juveniles found on beaches
that some active process of habitat selection is involved, perhaps mediated by the use of byssal thread flotation (Sigurdsson et al. 1976).
Like other Donax species, D. trunculus is relatively shallow burrowing, maintaining contact with the sand surface by means of separate inhalant and exhalant
siphons through which the respiratory and feeding current is drawn. The siphons are sensitive to vibrations and other disturbances that cause them to close and
withdraw but with little delay before extension and reopening occurs. Unlike many other Donax species (Ansell 1983), D. trunculus shows no directed tidal
migration. Its rapid burrowing movements, however, allow it to maintain its superficiallyburied position in the sand during disturbance by wave action. These
movements, combined with some surface movement by crawling or “leaping” probably result in redistribution of a proportion of individuals on each tide as is the case in
D. vittatus on Scottish beaches (Ansell 1994). In the longer term they lead to the size segregation of the population seen on Atlantic beaches. The characteristic
vertical and geographical distribution of Donax species on European beaches are related to different tolerances to environmental temperature extremes (Ansell et al.
1980a).
D. trunculus is reported as the host of a variety of parasites including trematodes (Pelseneer 1896, 1906, Lucas 1965, Mouëza 1975), gregarine sporozoans and
nematodes (Mouëza 1975), and a species of Rickettsia (Comps & Raimbault 1978). Where they infest the visceral mass and gonad, some parasites may cause
parasitic castration of the host (Pelseneer 1896) but none has been studied in any detail.
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Where it occurs in high densities, Donax trunculus is often the dominant filterfeeding macroinvertebrate on Atlantic beaches or in the shallow subtidal of the
Mediterranean Sea, feeding on suspended organic material, predominantly phytoplankton (Plante Cuny & Bodoy 1987). Its morphological adaptations to this role have
been well documented (Mouëza & Frenkiel 1974, 1976a,b, 1979, Mouëza 1976, Pichon et al. 1980). Beaches dominated by D. trunculus have not been extensively
studied at community level and no synthesis of its role in the food chain can be made. D. vittatus populations, which occupy a similar ecological niche on UK beaches,
are subject to predation by a variety of epibenthic marine predators, including drilling gastropods (naticids), crustaceans (shrimps, portunid and other crabs), and fishes
(particularly flatfishes). Predation may be lethal, resulting in the death and ingestion of the whole bivalve, but in many instances is nonlethal, often involving the loss of
only one siphon tip. This nonlethal (grazing) predation plays a major role in the nutrition of juvenile flatfishes like plaice, dab, and flounder that use sandy beaches as
nursery grounds (Ansell & Gibson 1990). It is likely that a similar suite of predators affect D. trunculus populations. George (1965) and NeubergerCywiak et al.
(1989, 1990), for example, report predation by naticid gastropods occurring in Mediterranean populations.
Population dynamics
Aspects of population dynamics of D. trunculus have been studied in Mediterranean (Bodoy & Massé 1979, Dowidar & El Nady 1984, NeubergerCywiak et al.
1990) and Atlantic (Degiovanni & Mouëza 1972, Mouëza 1972b, Guillou & Le Moal 1978, 1980, Ansell & Lagardère 1980, Bodoy 1982, Guillou 1982, Otero
1982, Fernandez et al. 1984, Bayed & Guillou 1985, Mazé & Laborda 1988, Le Moal 1993) populations. Several authors provide data on population densities of D.
trunculus in different areas (Atlantic: Degiovanni & Mouëza 1972, Mouëza 1972a, Guillou & Le Moal 1978, 1980, Ansell & Lagadère 1980, Bodoy 1982, Guillou
1982, Otero 1982, Fernandez et al. 1984, Mazé & Laborda 1988, Le Moal 1993; Mediterranean: NeubergerCywiak et al. 1990).
The maximum size reached by D. trunculus lies between 30mm and 44mm shell length. Maximum size varies among different populations, depending on both the
growth characteristics and on the extent of exploitation of the larger size groups (see later). Both the life span and the growth rate vary among populations from different
geographical areas, between different beaches in the same geographical area, and among different cohorts from the same beach. At the northern limit of the
geographical range in Brittany, up to six annual cohorts may be present in the population at any time, indicating a maximum life span in this area of more than six years
(Guillou & Le Moal 1980, Guillou 1982). Here, growth is restricted to the summer months and the shells bear well marked annual growth checks that provide a reliable
basis for growth measurements. Clear growth checks are also found in shells from further south on the French Atlantic coast (Ansell & Lagardère 1980). In Algeria, the
maximum sized individual found by Mouëza (1972b, 1975) was 34mm shell length. The population here showed numerous modal classes, but apparently few
individuals survive for more than 16–18 months. The life span is also short in other Mediterranean populations and at the southern end of the range in the Atlantic.
Growth and life span in other Atlantic populations lie between these two extremes, ranging between c. 2.90 and 3.25 (average ≈3.08, Table 2, p. 168).
Most authors have not attempted to estimate mortality or overall production. Ansell & Lagadère (1980) provided a “provisional” estimate of mortality and
production based on mean sizefrequency distributions (equivalent to “catch curves”) for two populations at Ile
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de Oleron, French Atlantic coast. These indicate a gradually increasing rate of mortality from year one to year four with few individuals surviving into their fifth year. On
the two beaches studied the mean estimated rate of production was similar, a lower mean population density on one being compensated for by a higher productivity per
individual resulting from a higher growth rate.
Fishery
The major commercial shellfisheries that exploit natural populations of intertidal or shallow subtidal soft sediments in European waters all centre on sheltered sand flats
or estuaries rather than on the more exposed beach habitats that form the subject of this review. D. trunculus is the closest analogue on European coasts to the beach
clams of the other geographical areas discussed and is included here for that reason. D. trunculus is exploited in most European countries although generally only as a
recreational fishery for individual consumption or as an artisanal commercial fishery supplying local markets. In some countries, there is a more extensive fishery; in Italy,
for example, D. trunculus is fished on the east coast from Tuscany to the southern part of the Napoli area, and on the Adriatic coast in the north (Friuli, Venezia Giulia)
and south (Puglia) (D.Del Piero pers. comm.). D. trunculus has been heavily exploited in some areas of Italy, notably the Tyrrenian Sea coast. Dowidar & ElNady
(1984) report that D. trunculus, known locally as “Om ElKholool”, is the most popular edible clam along the Mediterranean coast of Egypt where it is caught
commercially using a “special dredge, dragged by man or animal power” (Aleem 1969). There are no published official statistics for any D. trunculus fishery.
The techniques of harvesting D. trunculus vary from country to country and from region to region; they mostly involve methods of manual collection by individual
fishers. Otero
Figure 11 Hand dredge in use for collecting Donax trunculus in Portugal. (Photograph courtesy of Simon Grosset, photojournalist.)
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(1982) described the “small rasca”, a sicklelike device used by fishers in Galicia, Spain, to bring Donax to the sand surface. Along the coast of the Algarve, Portugal,
fishers use individual dredges drawn backwards in shallow water, with a harness worn round the waist, and rocked to and fro by means of a wooden post attached at
the front of the dredge (Fig. 11). On the French Mediterranean coast, a somewhat similar small dredge known as a “tellinière” is pulled by hand or, in some cases, by a
horse. Hydraulic dredges, normally employed in fisheries for Chamelea gallina (Veneridae) and Ensis minor (Solenaceae), were used in Italy to harvest Donax
trunculus but are now prohibited (D.Del Piero pers. comm.). In some areas local regulations have been introduced to help conserve the stocks of D. trunculus and
other exploited bivalves in shallow coastal waters; local regulations in Andalusia, Spain, for example, define a minimum size of 30mm shell length and a closed season
for the fishery extending from 1 April to 30 September (Dr Carmen Salas pers. comm.).
Africa
This section covers a single species that occurs on sandy beaches in South Africa and Namibia. Other species may support artisanal fisheries elsewhere in Africa but
this is the only species for which published information is available and it is probably also the only species that is, or has been, utilized commercially.
The white sand mussel Donax serra (Fig. 2i, p. 166)
Distribution
D. serra occurs around the southern African coastline from the Transkei coast of South Africa in the east through to the Kunene River at the border between Namibia
and Angola in the west (Fig. 1, p. 164). Its distribution thus covers three biogeographic provinces: the warm temperate south coast and the temperate Namaqua and
Namib upwelling regions along the west coast (Bally 1986, Donn 1990a). Intertidal distribution varies considerably between the warm temperate south coast province
and the cold temperate west coast provinces.
The intertidal distribution presents a distinct sizespecific zonation that differs between regions. In the east and south the species is entirely intertidal with adults being
centred on the midshore, although spat may settle in the surf zone and migrate upshore as they grow (McLachlan & Hanekom 1979, Donn 1990a, Lastra 1994). On
the west coast this reverses and juveniles occur in the intertidal with adults on the lower shore and in the surf zone (de Villiers 1975a, Bally 1983, Donn 1990a).
However, in northern Namibia intertidal distribution switches back to the south coast pattern, with adults in the intertidal (Bally 1986, Donn 1990a). This is probably
due to increasing water temperature north of the upwelling region in Namibia and parallels the seasonal shifts in distribution of Mesodesma mactroides in South
America.
Donax serra is not the only donacid inhabiting South African beaches; D. sordidus, a smaller tidal migrant species, occurs lower on the shore on the south coast.
There is no indication of any competition between the two species and distribution of the latter is centred on the surf zone, whereas D. serra is entirely intertidal except
for new recruits that settle in the surf zone.
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Distribution and zonation are not constant and this species, like most inhabitants of exposed beaches, undergoes migrations. In contrast to most other Donax
species, however, D. serra does not undergo tidal migrations, but rather shows small semilunar movements up and down the shore coupled to changes in beach water
tables during spring and neap tides (McLachlan et al. 1979, Donn et al. 1986). These movements have been recorded only in south coast stocks; west coast
populations do not appear to undertake any tidal or semilunar migrations. It has been suggested that the large size of D. serra and the low water temperatures on the
west coast make rapid burial difficult, precluding any migrations or intertidal existence, hence the subtidal adult stocks (Donn 1990a, Donn & Els 1990).
Another distribution pattern described for D. serra is a longshore sizespecific segregation around logspiral bays in the eastern Cape. Donn (1987) suggested that
spat recruit more abundantly near river mouths but are subsequently transported eastwards by longshore currents, thereby creating a size specific longshore zonation in
these populations: juveniles are most abundant near river mouths, whereas adults become more prolific eastwards. Dugan (1994) subsequently attempted to test for
longshore migration in adults by tagging, but failed to detect any significant movement. Furthermore, juvenile distribution was patchy with no clear size gradient as
postulated by Donn (1987).

Both south and west coast stocks of D. serra have a poorlydefined breeding cycle and exhibit extended spawning with, typically, more than one peak in the year. On
both coasts these peaks generally span the periods late summer and late winter. Sexes are separate and hermaphrodites have not been recorded; the sex ratio is 1:1
and they are external fertilizers (de Villiers 1975b, McLachlan & Hanekom 1979, van der Horst 1986). Females produce large numbers of small (45μm) eggs (van der
Horst 1986) but fecundity has not been quantified.
Seasonal changes in biochemical composition of adults have been linked to the reproductive cycle (McLachlan & Hanekom 1979). D. serra on the south coast
undergoes a substantial drop in tissue mass in winter and spring and a smaller drop in summer, corresponding to the two spawning seasons. The tissue comprises 66%
protein, 15% carbohydrate and 4% lipid.
A freeswimming planktotrophic larva develops through trochophore and veliger stages before settling as spat (Brown et al. 1989) but the duration of larval life is not
known. Recruitment is to the surf zone on the south coast but probably to the intertidal on the west coast. Settlement can occur throughout the year but may peak in
late summer and early autumn for the 2–3 mm length class on the south coast where spat appear to settle selectively in patches of fine sand in the surf zone (Lastra
1994). De Villiers (1975b) recorded peak recruitment to a length of 3–5mm during spring and summer, following the extended major spawning period starting in
winter.
D. serra, which is the largest member of the genus, is a successful suspension feeder and the dominant contributor to macrofaunal biomass on many South African
beaches (McLachlan et al. 1981, Donn 1986). The biology of the species has been reviewed by Brown et al. (1989). D. serra positions the tips of its siphons level
with the sand surface and takes in swash and surf water including phytoplankton and detrital material. The inhalant siphon has a fringe of tentacles to exclude sand
grains. It feeds in pulses and is capable of filtering a wide range of particles from 40μm down to bacteria, but extracts particles in the size range 4–7μm most efficiently
(StentonDozey 1989, Matthews et al. 1989).
The siphons bear ciliated tufts on their surfaces that may be primary receptors responding
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to light, tactile or chemosensory stimuli (Hodgson & Fielden 1984, Hodgson 1986). D. serra has a large bladeshaped foot and muscles plus the pedal haemocoelic
cavity are used to extend the foot and aid in burrowing (Brown et al. 1989). It is a rapid and powerful burrower (Trueman & Brown 1985) and burial time increases
with size and lowered temperatures (McLachlan & Young 1982, McLachlan et al. 1995). It also surfs in the swash, smaller individuals doing so more frequently. The
energetics of locomotion have been reviewed by Brown et al. (1989).
D. serra is relatively tolerant of heat stress, surviving for extended periods at temperatures of 25–30°C. Size classes that occur highest on the shore show the
greatest temperature tolerance on both coasts (Ansell & McLachlan 1980, StentonDozey 1989, StentonDozey & Brown 1994). D. serra is also moderately tolerant
of lowered salinities (McLachlan & Hanekom 1979) and could thus penetrate a short distance into estuaries if other conditions were suitable.
Considerable attention has been paid to respiration in D. serra and published information is summarized in Brown et al. (1989). The gills are not especially large in
relation to body size, oxygen uptake is proportional to body weight to the power 0.7–0.75 and animals display flat ratetemperature curves. They are oxyconformers,
uptake increasing in proportion to ambient partial pressures. Metabolism during withdrawal into the shell under unfavourable conditions is anaerobic, the animal
incurring an oxygen debt. An energy budget compiled for this species indicates that scope for growth is greatest when it is fed on algal rations and lower when fed on
detritus (Brown et al. 1989, Matthews et al. 1989). No parasites or commensals have been recorded on D. serra. With high fecundity, rapid growth and unspecialized
feeding, D. serra is clearly a typical rstrategist species
D. serra is the dominant suspension feeder among the benthos on both south and west coasts of southern Africa. Its standing stock can run into kilograms of dry
flesh mass per linear metre of beach and it constitutes up to 98% of intertidal biomass. Its position in the ecosystem is best understood in the eastern Cape sector of the
south coast where food webs, trophic relations, energy flow and nutrient cycling in beach and surfzone ecosystems have been well studied. Trophic dynamics of the
beach and surfzone ecosystem in this area have been summarized in reviews by McLachlan & Bate (1984), Cockcroft & McLachlan (1993) and Hey mans &
McLachlan (1996).
This beach and surfzone ecosystem is fuelled by intense accumulations of surf diatoms, Anaulus australis. Within the macrofaunal trophic assemblage, zooplankton
and benthos constitute approximately equal biomass and D. serra makes up 70% of the latter biomass. It has a P:B ratio of about 0.6yr−1 (McLachlan & Hanekom
1979, Donn 1986, Schoeman 1996). Production by this species constitutes about 10% of all production in the macrofauna assemblage and is taken by swimming crabs
Ovalipes trimaculatus (30%), birds, especially oystercatchers and gulls (40%), and a variety of benthic feeding fishes, including batoids (30%) (du Preez 1984,
McLachlan et al. 1980, du Preez et al. 1990). Predation is both lethal, for example sandsharks preying on juveniles in the surf zone and skates, oystercatchers and gulls
taking adults in the intertidal, and sublethal, with sanderlings nipping siphons in the intertidal and some fishes doing this in the subtidal. Damaged siphons can be
regenerated within a few weeks (Hodgson 1986).
As well as its contribution to energy flow, Donax serra is also a regenerator of nutrients, principally ammonia, which represents a significant source of nitrogen to
phytoplankton (Prosch & McLachlan 1984, Cockcroft & McLachlan 1993). Its shells contribute to the calcium carbonate fraction on these beaches.
The above scenario applies to the eastern Cape sector of the south coast where exposed inter
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mediate to dissipative beaches are characterized by high surfdiatom production. Elsewhere the role of white sand mussels may differ; on intermediate to reflective
beaches it is much less abundant or absent. On the west coast, surf zones are characterized by upwelling, high detrital inputs from kelp beds and the absence of surf
diatoms. The great biomass of sand mussels in the surf zones of the more dissipative beach types suggest that they must play an important role as consumers of water
column particulates and as food for fishes and crabs. The consumption of adults by birds is restricted to the south coast where adult white sand mussels occur
intertidally.
Population dynamics
The population dynamics of D. serra have been examined in two areas, on the west coast north of Cape Town and on the eastern sector of the south coast near Port
Elizabeth. The west coast studies (de Villiers 1975a) could only clearly distinguish two age groups on the beach because slowing of growth with age resulted in a
unimodal frequency band of adults and this was compounded by the migration of adults into the subtidal below the limits of sampling. Recent stock assessments (Tarr
1994) at four west coast sites indicated mean population sizes on the beach and shallow subtidal of 150–4272 per linear metre (m−1), with a maximum of 7884m−1,
and populations up to 4.3 million km−1 shoreline. On the south coast, populations have been studied by McLachlan & Hanekom (1979), Donn (1986) and Schoeman
(1994, 1996). Most work has concentrated on two exposed beaches (24km and 40km long) where abundances are high, ranging from 1000–7000m−1 (mean
2300m−1) and shellfree dry biomass values up to 7kgm−1 (mean 1600gm−1) have been recorded. Abundance shows interannual fluctuations, mainly due to varying
recruitment, adult stocks being remarkably stable.
Growth is rapid, to 32–35mm in the first year in the intertidal, and to 48–50mm in the second year, on the south coast (McLachlan & Hanekom 1979, Donn 1986,
Schoeman 1996). Thereafter growth rate decreases to a size of 67mm (maximum 80mm) after five or more years (Schoeman 1996). Empirical (McLachlan &
Hanekom 1979), Gompertz (Donn 1986) and von Bertalanffy seasonally oscillating (Schoeman 1996) growth curves have been estimated, all in close agreement: K
ranged 0.39–0.73yr−1, and L∞ 66–68mm. An approximate value of the growth index was 3.57 (Table 2, p. 168).
On the west coast growth is slower but continues until a larger size is attained. De Villiers (1975a) was able to obtain growth ring readings of sufficient reliability to
compile a growth curve that corresponded well with curves from lengthfrequency analyses. De Villiers (1975a) estimated an empirical growth curve with lengths of
22mm, 43mm, 58mm, 64mm and 68mm after their first five years on the beach and sexual maturity at 37–54 mm or 1.5–2.5 years. West coast specimens can grow to
80mm shell length. This species does not usually have distinct and usable growth rings on the south coast.
There was remarkable consistency in these three studies and P:B ratios were found to be about 0.6yr−1 and total mortality 0.5–0.85yr−1 with fishery mortality
(F=0.07yr−1) negligible (Schoeman 1996). Comparable production and mortality data are not yet available for the west coast. Sexual maturity is reached at a length of
about 44mm on average (range 38–49mm), this being at an age of about 2yr after settlement or 1.5yr in the intertidal (McLachlan & Hanekom 1979, Schoeman
1996).
Donn (1990b) analyzed morphometric differences between west and south coast populations, finding the slightly smaller intertidal south coast specimens to have
heavier valves, greater density and more wedged shapes than their colder water west coast counterparts. He suggested that these differences aided better movement
and rapid burial in the swash in the case of south coast specimens.
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Mass mortalities are a common phenomenon on the west coast and de Villiers (1975a) described one major mortality at Elands Bay. This was caused by red tides in
the form of blooms of the dinoflagellate Gonyaulax grindleyi. This effect was greatest at lower tide levels and virtually all adult individuals perished; recovery was not
evident within six years. Such red tides are a common occurrence on the west coast but only occasionally cause mass mortalities on the scale described by de Villiers
(1975a). On the south coast minor mortalities have been recorded where many dead animals littered the drift line. It is not known what caused such mortalities but they
generally affect less than 5 % of the population at any one time (A.McLachlan unpubl. data).
Fishery
Stone age peoples around southern Africa subsisted on sand mussels for millennia and their shells characterize middens along the coast. In recent times mussels have
been collected in recreational and commercial fisheries for use as bait and food, but mainly bait. The west coast stocks have been subjected to both forms of fishery but
only recreational fisheries exist on the south coast. A size limit of 38mm shell width (58mm length) has long been in force. Statistics of licensed landings for commercial
collecting, recorded since 1966, show a clear decline (Fig. 12) coupled to both decreasing demand and reductions in annual quotas. All licences were revoked in 1988
and the issue of new licences has been limited pending the outcome of ongoing stock assessments (Tarr 1994). The problem of toxins accumulating in the tissues during
red tides has inhibited the development of this industry on west coast beaches and landings are used primarily for bait. Although little further information is available on
this fishery it is clear that these stocks are lightly utilized and could sustain a much greater harvest if it were not for the inhibiting effects of poisoning by red tides.
On the south coast, intertidal adults are widely collected in a recreational fishery. Although some interest has been expressed in establishing a small commercial
operation near Port Elizabeth, there has been strong public opposition to what is seen as a danger to the wilderness quality of the beaches and no formal applications
for permits for such a fishery have been submitted to date. Schoeman (1996) and Sims (1996) have monitored the recrea
Figure 12 Annual commercial landings of Donax serra for bait. (After Tarr 1994.)
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tional fishery on the beach supporting greatest sand mussel populations and found that collectors remove <2% of the population (<2% of biomass) per year. However,
many collectors exceeded their quota of 50 mussels per day and collected animals below the size limit. The infringement of size limits is largely because the size limit of
58mm shell length, 14mm> than size at first maturity (Table 2, p. 168), is inappropriate for the south coast where adults are smaller than on the west coast and <10%
of the population is above this minimum limit. D. serra appears to be underexploited on the south coast and there is an as yet untapped potential for a commercial
fishery. An ongoing study of possible harvesting impacts and a costbenefit analysis of the feasibility of a commercial fishery for D. serra in St Francis Bay on the south
coast have indicated that 10% of clams are damaged during harvesting, even using traditional techniques (spades), but that no ecological damage to the beach could be
detected. These results indicate that a small commercial fishery is indeed viable (Schoeman 1996, Sims pers comm.).
There is a fundamental difference between the south and west coast stocks of D. serra in terms of fishery management. The intertidal situation of adults on the south
coast makes them readily accessible to collecting and the absence of red tides allows utilization for food and bait. Careful control and management is thus needed on
the south coast and quotas and collection methods (hand only) should be enforced. However, minimum legal size on the south coast could be reduced to about 45mm
shell length, since this is above size at maturity; it could reduce incidental mortality and is more appropriate for a population that has a smaller individual size than on the
west coast. On the west coast, however, the subtidal distribution of most of the adult stock renders them relatively inaccessible, red tides prevent their use for food at
certain times and the considerable extent of the coast and low human population densities also limit impact. It is therefore doubtful whether any management or control
is needed beyond quotas and size limits.
Asia
Numerous species of bivalve are harvested and utilized as food in tropical and subtropical countries around the IndoPacific region, but most of these fall outside the
scope of this review as they are either subtidal in distribution, or from intertidal sand flats or estuaries. Sand beaches in this region commonly support populations of
mostly smallsized donacid, mesodesmatid and venerid bivalves; only the few larger species are exploited in some areas by mainly artisanal fisheries, although the
commercial potential of other species has been explored in some countries (Mwaidseje 1982, Parulekar et al. 1984).
The wedge clams Donax cuneatus and D. faba (Fig. 2j, k, p. 166)
Distribution
D. cuneatus is distributed on tropical coasts from east Africa, throughout the IndoPacific region to Japan and Australia, and eastward in the Pacific Islands to Fiji (Fig.
1, p. 164). Typically it is found on intermediate or dissipative beaches (Ansell 1985). The closely related species D. faba has a similar geographical distribution, but is
typically found on more reflective beaches of relatively coarse sand (Smith 1975, Ansell 1985). Populations of both species undergo some tidal migration (Smith 1975,
Ansell 1985) and may also show some
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vertical separation in the distribution of juveniles and adults on the beach (Alagarswami 1966). The two species may sometimes have been confused in the past as both
are variable in colouration and shell shape, although there are clear and consistent taxonomic differences (Ansell 1985).

As in other Donax species, in both D. cuneatus and D. faba the sexes are separate. The sex ratio does not differ significantly from 1:1 in D. faba (Alagarswami
1966), but Nagabhushanam & Talikhedkar (1977a) found a 1.1:0.9 ratio in favour of females in D. cuneatus. In both species, first maturity is reached in less than a
year from settlement; in D. cuneatus at about 10–11 months and a length of 12–13mm (Nayar 1954, Talikhedkar et al. 1976) and mD. faba at 6–7 months and 10–
11 mm (Alagarswami 1966) (Table 2). Where they have been studied, the two species show very similar reproductive cycles, with distinct seasonal peaks in spawning
that are reflected in seasonal peaks in settlement and recruitment to the beach populations. In India, where the two species have been most studied, the seasonality in
the reproductive cycle is related to changes in temperature and particularly salinity brought about by the seasonal monsoon cycle. In D. cuneatus, at Ratnagiri on the
east coast of India, development of the gonads began in March and by July the whole population was in a fully ripe condition that was maintained from July to
September; spawning extended from October to January, reaching a peak in November–December (Nagabhushanam & Talikhedkar 1977a). Differences in the timing
and duration of development and spawning stages of D. cuneatus, however, have been reported for other localities (Nayar 1954, Rao 1967, Talikhedkar et al. 1976).
In D. faba, from the Gulf of Mannar on the southeast coast of India, development of the gonads took place between August and late October, by which time all
individuals in the population were ripe; spawning began in November and most individuals were spent by the end of that month—although some spawning continued
into December; a further development of the gonad then took place through the period January to April with further spawning from March through June (Alagarswami
1966). Changes in tissue weight and the biochemical composition of the tissues are related to the gonadal cycle (Alagarswami 1966, Nagabhushanam & Talikhedrar
1977b).
D. cuneatus and D. faba are both rapid burrowers. In D. cuneatus the BRI increased from <5 to >15 over the range 15–26.5°C (Ansell 1985) indicating that it
burrows more rapidly than any of the other beach clams reviewed here for which this index has been determined (McLachlan et al. 1996). D. faba had a lower
burrowing rate index at 25°C than D. cuneatus (Ansell 1985).
Both species can develop large populations where they occur on productive beaches, becoming dominant benthic filter feeders (Harkantra & Parulekar 1986),
Smith (1975) recorded densities of up to >2000m−2 for D. faba on Tanzanian beaches. Hughes (1966) reported that D. faba are heavily predated by the ocypodid
crab Ocypode ceratophthalma on beaches in Mozambique and Smith (1975) showed that this crab selectively attacked different colour morphs in populations on the
coast of Tanzania, East Africa. Smith also reported predation by the sanderling, Crocethia alba in the same area.
Population dynamics
Relatively little information is available on the population dynamics of either D. cuneatus or D. faba over most of their geographical range. Nayar (1954) and
Talikhedra et al. (1976)
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studied the growth rate of D. cuneatus, and Alagarswami (1966) that of D. faba, in India. Shell growth is rapid, more or less constant and steady during the initial
growth period of about one year, but then slows (Alagarswami 1966) or stops after sexual maturity is reached, although there may be further short periods of renewed
growth thereafter (Nayar 1954). D. cuneatus can potentially reach a larger size than D. faba, although Alagarswami (1966) recorded a maximum length of 27mm for
D. faba, and Nayar (1954) only about 23mm for D. cuneatus (Table 2, p. 168), however, in both species the maximum shell length reached can vary greatly among
populations in different localities. The maximum life span probably does not exceed 3yr (Table 2).
Fishery
Wedge clams in India are exploited by the coastal fishing communities, but do not form the basis of a large commercial fishery. Both Nayar (1954) and
Nagabhushanam & Talekhedkar (1977b) stated that D. cuneatus is exploited mainly at times when general fishery conditions are poor and other shellfish difficult to
obtain, for example particularly during the monsoon season when the sea is rough. Elsewhere, as in the Phillipines (Talavero & Faustino 1933), artisanal fisheries exploit
local populations of D. faba. In some parts of India (Chari & Unny 1947) and in the Phillipines (Talavero & Faustino 1933) the shells have been used in the production
of quicklime.
Australia
Only a single clam, the donacid Donax deltoides, is widely utilized on sandy beaches in Australia. It has not been well studied and the limited information available is
summarized here. Hopefully this will encourage further study since this appears to be the only species of beach clam in Australia with commercial potential.
The pipi or Goolwa cockle, Donax deltoides (Fig. 2l, p. 166)
Distribution
D. deltoides occurs along the south and east coasts of Australia from the Adelaide area up to southeastern Queensland, and including Tasmania (Fig. 1, p. 164).
Juvenile clams are located in the intertidal and adults on the lower shore and in the surf zone, but their distribution is probably centred on the low tide swash zone area
(Dakin & Bennett 1987, Saenger & Keyte 1990, McLachlan et al. 1996). It is possible that juveniles undergo tidal migrations but unlikely that the adults do so. In
many respects, including appearance and distribution, this species is very similar to D. serra in South Africa.

Almost nothing is known of the reproductive biology of D. deltoides. In South Australia spawning can occur throughout the year but is most marked in spring. Sexes
are separate and pipi are probably broadcast spawners. There is a 6 to 8 week larval phase before settlement (Saenger & Keyte 1990).
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D. deltoides develops large populations on high energy dissipative beaches where surf diatom blooms occur, e.g. Goolwa (McLachlan & Hesp 1984). Here they
are the dominant benthic filter feeders, for example making up 73% of macrofauna numbers and 85% of biomass on Goolwa beach (McLachlan et al. 1996), and thus
occupy a key position in the beach and surfzone food web. Their predators, besides man, include beachworms (Onuphis sp., Dakin & Bennett 1987), seabirds and
fishes.
Population dynamics
Donax deltoides is said to exhibit resurgent populations but supportive data are lacking. River discharge and calm periods without vigorous wave action are
considered possible causes of widespread mortalities (Saenger & Keyte 1990), but no studies have been published on the population dynamics. There are a handful of
estimates of density, with values ranging up to 345m−2 (juvenile densities can exceed 1000m−2). Growth is rapid; clams reach 36mm shell length and sexual maturity in
13 months. Growth continues to a maximum length of about 60mm in 3.5yr (Saenger & Keyte 1990, unpublished data from the South Australian Fisheries
Department). Only one empirical growth curve has been determined for South Australia stocks of D. deltoides (Saenger & Keyte 1990), which rendered preliminary
estimates of the growth parameters L∞ and K close to 56mm and 1.59yr−1, respectively, giving a value of the growth rate index of 3.70 (Table 2, p. 168).
No information is available on recruitment or mortality for D. deltoides.
Fishery
In the past, D. deltoides was a staple item in the diets of coastal aboriginal populations (Godfrey 1988). The fishery for pipi in Australia consists of both recreational
collecting, mostly by anglers for bait, and commercial harvest for eating. This is undertaken in South Australia and New South Wales and possibly also Victoria. No
reports are available on landings. However, Saenger & Keyte (1990) failed to detect any significant reduction in pipi densities at beaches where they were subject to
commercial exploitation in comparison with unexploited populations in northern New South Wales. There is no management of this resource other than size and quota
limits: 50 clams of any size per person per day in New South Wales, a 5–1 bucket of clams of any size per person per day in Victoria and any number of clams above
35mm length in South Australia.
New Zealand
Three species of clams of the genus Paphies have supported traditional, recreational and commercial fisheries on New Zealand sand beaches. The taxonomic history of
these species is complicated; there have been more than 30 names applied to P. subtriangulata since 1828 (Beu & De RooijSchuiling 1982). The recent division of
the tuatua species into P. subtriangulata and P. donacina by Beu & De RooijSchuiling (1982) has been supported by electrophoretic and morphometric studies by
Richardson et al. (1982) and Smith et al. (1989).
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The toheroa Paphies ventricosa (Fig. 2m, p. 166)
Distribution
P. ventricosa is endemic to New Zealand and populations occur at scattered locations throughout the country (Fig. 1, p. 164). Historically, the largest populations
were found on three, long, northwestern coast beaches on North Island: Ninety Mile Beach (89km), Dargaville Beach (72km), and Muriwai Beach (48km). Elsewhere
on North Island, substantial populations have also been found at Mitimiti and on the Wellington west coast beaches, particularly Hokio and Waitarere (20km total), and
small populations have been reported from the northeastern coast. On South Island, significant stocks are found on the southern coast at Oreti Beach (17km) and Te
Waewae Bay (11km), and small populations have occasionally been reported on the southeastern coast (Redfearn 1974).
Populations of P. ventricosa are found intertidally on wide, fine sand beaches (Redfearn 1974). The largest populations occurred on dissipative beaches with
prevailing onshore winds and regular surfzone diatom blooms on the west coast of North Island (Cassie 1955). Large P. ventricosa at these sites were aggregated in
dense beds. Redfearn (1974) observed adult aggregations and higher densities of recruitment in “bays” adjacent to rip currents. Street (1971) reported that populations
at Te Waewae Bay did not occur in distinct aggregations.
Intertidal zonation of P. ventricosa is associated with size (Redfearn 1974, Street 1971). Large clams (>75mm shell length) are generally aggregated in beds from
the mid to low intertidal. Smaller clams are usually found on the upper beach or in separate singleaged beds in the mid intertidal, but may also occur in a layer between
large clams and the sand surface. Dense bands of newly recruited clams have been found just below the high tide line (Redfearn 1974). Although Cassie (1955)
proposed that subtidal stocks must exist to explain fluctuations in stock estimates and missing small cohorts in size frequency distributions, no subtidal stocks have been
documented despite numerous attempts (Street 1971). Toheroa, P. ventricosa, occur from just beneath the sand surface to depths of 20cm (Redfearn 1974). In good
conditions, toheroa can be located by siphon holes visible on the sand surface at low tide (Cassie 1955).

Reproduction of P. ventricosa is by broadcast spawning, and fertilization is external. Sexes are separate and have a ratio of close to 1:1 (Redfearn 1974). Sexual
maturity occurs at small sizes. Redfearn (1974) reported that some juveniles had fully developed gonads at 10mm shell length, the majority of individuals matured in
their first year, and all individuals were mature by 15 months (47mm). The main spawning season on Dargaville beaches occurred from September to February, but
individuals with ripe gonads were found throughout the year (Redfearn 1974). Fecundity varies with female size; 15–22 million eggs, 60–66μm in diameter, may be
produced in a single spawning in the laboratory (Redfearn 1982).
Larvae of P. ventricosa have been reared successfully in the laboratory with development being completed within 22 days at 25°C (Redfearn 1982). Settlement
occurred in the laboratory at lengths of 270–300μm (Redfearn 1982).
The siphons of P. ventricosa are the longest in the genus in New Zealand, and allow toheroa to feed while deeply buried. The siphons are held close to the level of
the sand surface when feeding. Adult P. ventricosa dig rapidly when disturbed, reaching depths of
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200mm (Redfearn 1974). For juvenile P. ventricosa, the average burrowing rate index (BRI) was 4 at 15°C (J.E.Dugan & D.M.Hubbard unpubl. data), a value
intermediate between those of juvenile P. subtriangulata (BRI=7) and juvenile P. donacina (BRI=3) (McLachlan et al. 1995). Redfearn (1974) observed small
clams, and occasionally larger ones, surfing in the swash. Aggregations of toheroa may move during disturbance events, and have been reported moving seaward by as
much as 30m in a day in the absence of noticeable disturbance (Redfearn 1974), but some beds remain stationary for years (Rapson 1952). There is no evidence of
tidal migration in P. ventricosa.
The condition of P. ventricosa varied seasonally at Dargaville and was lowest in the summer during the peak spawning season. Meat volume as a percentage of
shell cavity volume ranged from maximum values of >70% in August and September to <50% in January and February (Redfearn 1974).
A variety of predators including birds, crabs and fish prey on P. ventricosa. Southern blackbacked gulls, redbilled gulls, South Island pied oystercatchers and
variable oystercatchers prey on toheroa during low tide. Southern blackbacked gulls have been reported taking clams up to 120mm in length (Redfearn 1974). The
effects of paddle crab, Ovalipes catharus, predation at high tides is probably significant only for small size classes of clams (Wear & Haddon 1987). Results of
laboratory experiments suggest that individual Paphies ventricosa attain some refuge from Ovalipes catharus predation with increasing depth of burial, and mortality
is reduced in dense aggregations. Paphies ventricosa are also preyed on at high tide by fish including snapper, Chrysophrys auratus (Rapson 1952).
Population dynamics
The earliest stock estimates available for Paphies ventricosa are from the 1930s, and were made after decades of commercial harvest and the collapse of at least one
major stock at Muriwai in 1928 (Redfearn 1974). Stock estimates from most of the important fishery areas for P. ventricosa have declined to about 10% of their
highest levels in the last few decades (Fig. 13). Maximum densities reported for the six major fishery areas are: 340ind.m−1 at Ninety Mile Beach, 420ind.m−1 at
Dargaville, 310ind.m−1 at Muriwai, 125ind.m−1 at Hokio and Waitarere, 90ind.m−1 at Te Waewae Bay, and 120ind.m−1 at Oreti Beach (harvestable clams only). The
only population that appears to have been relatively stable in recent decades is at Oreti Beach, South Island, which had an estimated density of 74 clams (>80mm)m−1
in 1990 (Millar & Olsen, 1995). Population estimates available for P. ventricosa have large confidence limits, and stock estimates fluctuate considerably from year to
year. Most of the available data on toheroa populations were obtained in efforts to assess harvestable stocks, and are likely to be biased against smaller clams both
because of the sampling methods and locations (Street 1971, Redfearn 1974).
P. ventricosa are longlived (>1Oyr) and have indeterminate growth. They have the fastest growth rate ( ≈3.83: Table 2, p. 168) and attain the largest size
(160mm, >470g) of the Paphies species in New Zealand and reach legal size for harvesting (100mm shell length, 120g) in 4 to 5 years on North Island beaches
(Cassie 1955), and 7 to 9 years on South Island beaches (McKinnon & Olsen 1994) (Table 2). Redfearn (1974) reported similar shell growth estimates using three
methods: length frequency analysis, size at age from shell rings, and markrecapture studies.
Size frequency distributions of P. ventricosa are not generally characterized by large numbers of youngoftheyear animals and decreasing numbers of older year
classes (Cassie 1955, Redfearn 1974, McKinnon & Olsen 1994). Samples of P. ventricosa populations gen
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Figure 13 Population estimates for Paphies ventricosa. North Island (top 4 beaches), South Island (bottom 2 beaches). Some South
Island estimates are for harvestable stocks only (>75mm 1971–78, >100mm 1979–90). Data are from Cassie 1955, Greenway
1969, Redfearn 1974, McKinnon & Olsen 1994, Millar & Olsen 1995, and unpublished MAF fisheries reports. Not all beaches
were surveyed each year and effort varied between surveys.
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erally have relatively few or no recruits, one or two small cohorts of intermediate size and a mode of larger animals presumably composed of several year classes
(Redfearn 1974, McKinnon & Olsen 1994).
Recruitment of P. ventricosa appears to be sporadic (Street 1971, Redfearn 1974). Cohorts from strong recruitment years do not necessarily survive to make
significant contributions to adult stocks (Greenway 1969, Street 1971, Redfearn 1974). Peak recruitment to the beach at Dargaville was in the summer, between
November and March (Redfearn 1974), but newly settled individuals were found as late as August. Redfearn (1974) observed small clams (2mm) being transported
up the beach by swash at all tidal phases, and concluded that this led to the formation of a concentrated band of recruits in the upper intertidal zone.
Population structures of P. ventricosa have changed on some beaches with the decline in harvestable stocks. For example, large clams (>100mm) have virtually
disappeared from Ninety Mile, Dargaville, and Muriwai beaches. In 1986, samples from these sites had substantial modes of clams between 40 and 50mm and <1%
clams longer than 100mm (MAF unpubl. data). Redfearn (1974) estimated mortality rates of P. ventricosa of about 50% per year, but was unable to determine what
proportion was due to harvesting.
Fishery
Toheroa have been subject to artisanal, recreational and commercial fisheries. Maori people harvested toheroa before the settlement of New Zealand by Europeans,
and middens contain large deposits of P. ventricosa. Commercial harvest of toheroa occurred from about 1900 until 1969 (Stace 1991). Canning started on North
Island in Dargaville in the 1890s (Redfearn 1974). Other canneries operated at various times at Waipapakauri at Ninety Mile Beach (1923–45), Muriwai (1946), the
Wellington beaches, and on South Island at Te Waewae Bay. The last canning licence was issued in 1969, and the last cannery was closed in 1971 (Stace 1991).
Longterm landings data are not available, but the production of toheroa products by canneries was summarized by Redfearn (1974). Maximum annual commercial
production of P. ventricosa peaked at 77t canned in 1940 (Fig. 14). A conversion of production data to landings is approximately 20 clams per kg canned (using data
from Dargaville, Redfearn 1974). This ratio varied due to seasonal and annual differences in the condition of the clams, size classes harvested, and the proportion of
clams in the canned product (e.g. as soup or whole clams).
Toheroa were harvested commercially with flat pronged potato forks at low tide. Rapson (1952) reports that at Dargaville beaches, 1 to 2m wide trenches were
dug through dense beds, and the fishers deliberately left areas with high densities of clams along the outside of the trenches. Some canneries operated seasonally, and
harvested when clams were at peak condition (Redfearn 1974). After 1962, quotas for the commercial harvest of P. ventricosa were based on annual surveys
(Redfearn 1974). Live toheroa were difficult to market. Mestayer (1921) attributed the short shelf life of live P. ventricosa to the valve gape of market size clams.
Canned whole clams and toheroa soup were produced for domestic sales and exports (Cassie 1955). Toheroa soup was typically bright green due to high
concentrations of phytoplankton in the guts of the clams.
No longterm data on recreational harvests are available, but recent estimates indicate that they can make substantial impacts on stocks in a short period. Estimated
recreational landings of P. ventricosa on Dargaville beach from 1966 to 1971 were between 160000 and 3700000 clams per season (Redfearn 1974). In 1974,
52000 toheroa (38% of the stock) were harvested in 11 days from an estimated harvestable population of 130000–136000 at Hokio
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Figure 14 Commercial production (tonnes canned) of Paphies ventricosa from Ninety Mile Beach, Dargaville and Muriwai, 1930 to
1970. (From Redfearn, 1974).
on the Wellington west coast. In 1975, approximately 1 million clams (52% of the stock) were taken at Dargaville in a 14 day opening, leaving an estimated 926000
remaining on the beach. Legal recreational take was 76% of the stock in fisheries openings of one week duration at Wellington west coast beaches in 1978 (127000 of
167000 takeable clams from three beaches) and 1979 (60000 of 79000 from four beaches). An estimated 56500 P. ventricosa (6% of stock) were harvested in 9h
at Oreti Beach in 1990 (McKinnon & Olsen 1994).
Recreational harvest of P. ventricosa was unregulated in 1921 (Mestayer 1921), and since then regulations on season, bag limit, size and methods have been
increasingly restrictive as stocks declined in most areas. In 1932, a 2month closed season, 50 clam per day bag limit, and minimum shell length of 76mm were
imposed. A bag limit for Maori fishers of 80 clams per day was set in 1941. In 1953, harvest was restricted to a 2month open season, and bag limits were reduced to
20 clams per fisher per day. The use of digging implements for P. ventricosa collection was prohibited in 1962. All North Island beaches were closed to toheroa
harvest for the year in 1967. Open seasons were reduced to two weeks in 1972, 7 days in 1978, and 5 days in 1980. Minimum shell length for harvesting was
increased to 100mm in 1979. P. ventricosa has been under total protection from all harvesting on North Island since 1980 (Haddon 1988).
The only two fishery openings since 1981 were single days at Oreti Beach on South Island (an 11h opening in 1990 and a 9h opening in 1993). Restrictions for the
one day southern fishery recreational openings included: (a) a minimum stock projection of at least 1000000 legal sized animals remaining on the beach after the season
at Oreti Beach and 450000 at Te Waewae Bay, (b) a closed reserve area at the south end of Oreti Beach, (c) a limit of five clams per person, and (d) no vehicles on
the beach. The impact of illegal take on stocks of P. ventricosa is unknown, but may be significant.
The continued decline in toheroa stocks have been attributed to many factors over several decades: increased harvesting rates, illegal harvesting, changes in currents
and phytoplankton production, mass mortalities, liquefaction of the sand by vehicular traffic, and sedi
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ment changes (Te Waewae Bay). Management efforts, including restrictions on harvesting season, size, bag limit, methods of digging, and closure of some fishery areas
for over 20yr, have failed to restore stocks of this highly prized species to levels capable of sustaining fisheries.
The tuatua Paphies subtriangulata (Fig. 2n, p. 166)
Distribution
P. subtriangulata is endemic to New Zealand, and occurs on beaches on North Island, South Island, Stewart Island and in the Chatham Islands (Fig. 1, p. 164).
Electrophoretic studies by Smith et al. (1989) suggest that there are three or four geographic groups of P. subtriangulata around New Zealand. Of these, the
Chatham Island group is most isolated and has reduced heterozygosity. Shell morphology varies between beaches (J.E.Dugan & D.M. Hubbard unpubl. data).
Populations of P. subtriangulata occur across a remarkable range of beach types from high energy dissipative beaches to sheltered reflective beaches (J.E.Dugan
& D.M. Hubbard unpubl. data). Adult P. subtriangulata generally occur from the low intertidal to shallow subtidal zones, but are occasionally found in dense beds in
the mid intertidal (Grant 1994). Recruitment occurs in the intertidal zone, and individuals are reported to move to lower levels over time (Greenway 1969, Grant 1994).

P. subtriangulata are broadcast spawners with separate sexes. The sex ratio is not significantly different from 1:1 (Grant 1994). Greenway (1981) reported that low
condition index values for P. subtriangulata from Dargaville between October and December occurred during the spawning season. Grant (1994) found two major
spawning periods at Little Omaha Bay, North Island, (October/November and February/March) and no resting period. Spawning was observed at high tide by Grant
(1994). During spawning events clams extended their siphons beyond the usual distance from the bottom and expelled visible streams of gametes. Some individuals
spawned for over 20minutes, and some aggregations exhibited spawning activity for up to 2h (Grant 1994). Female clams produced eggs that were about 60μm in
diameter in the laboratory (Redfearn 1987), and larvae were reared successfully. Larvae settled at a size of 230µm after 17 days at 20°C (Redfearn 1987).
P. subtriangulata are suspension feeders with short siphons relative to P. ventricosa. In a study of Wellington west coast beaches, algae and hydroids present on
the shells of P. subtriangulata were taken as evidence that they occupy shallower depths in the sediment than P. ventricosa and P. donacina (Richardson et al.
1982). P. subtriangulata have a burrowing rate typical of beach clams. McLachlan et al. (1995) reported that the BRI for juvenile clams was 7 at 20°C. Juvenile
clams have been observed surfing in the swash zone with feet and siphons extended (J.E.Dugan & D.M.Hubbard unpubl. data). There is no evidence of tidal migration
in P. subtriangulata. Predators on P. subtriangulata include birds, crabs and fish. P. subtriangulata form a considerable part of the diet of paddle crabs, Ovalipes
catharus (Wear & Haddon 1987), but clams >40mm shell length may have some refuge in size. Sublethal predation in the form of siphon nipping by O. catharus has
been observed during high tides (Grant 1994).
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Population dynamics
Few studies have investigated the population dynamics of Paphies subtriangulata. Population structure of P. subtriangulata in the low intertidal and shallow subtidal
was stable over 10 months (Grant 1994). Most individuals were between 30–65mm shell length, and the mean length of the dominant cohort was about 55mm (Grant
1994). Size structure and abundance of P. subtriangulata varied with beach morphodynamic type at 12 North Island sites. Clams from reflective beaches had larger
average sizes than those from more dissipative beaches (J. E.Dugan & D.M.Hubbard unpubl. data).
Abundance of P. subtriangulata has been estimated for intertidal (Grant 1994, J.E. Dugan & D.M.Hubbard unpubl. data) and shallow subtidal areas (Cranfield et
al. 1993, Grant 1994). Variability in abundance associated with beach type and harvesting pressure may affect the proportions of clams found in the two zones.
Densities of P. subtriangulata were as high as 1400m−2 in the midintertidal at Ocean Beach in May 1992 (Grant 1994). Low intertidal and shallow subtidal densities
averaged from 10 to 20ind.m−2 with maximum values of 55m−2 in adult aggregations at Little Omaha Bay (Grant 1994). Estimates of intertidal abundance at 12 North
Island beaches were correlated with Dean’s parameter and ranged from 25m−1 on a reflective beach to 495m−1 on a dissipative beach (J.E.Dugan & D.M.Hubbard
unpubl. data). Much higher densities of P. subtriangulata have occasionally been reported: Grant (1994) estimated up to 6000m−1 at Ocean Beach in 1992.
Estimated densities of larger P. subtriangulata from hydraulic dredge sampling in the subtidal ranged from 0–136m−1 (Cranfield et al. 1994), and biomass estimates
were 0–1.3kgm−1. Intertidal biomass estimates from 12 North Island sites ranged 0.2–7.8 kgm−1 and were not correlated with measures of beach state (J.E.Dugan &
D.M.Hubbard unpubl. data).
P. subtriangulata attains a smaller maximum size and has a slower growth rate ( =3.30; Table 2, p. 168) than its New Zealand congeners, P. ventricosa and P.
donacina. Grant (1994) estimated that clams reached 40mm shell length in 2yr, 50mm in 3yr and have a maximum age of more than 5yr. Large individuals reach
lengths of >80mm, and weights of >90g. Greenway (1981) estimated that P. subtriangulata took 3 yr to grow to a size suitable for commercial harvesting.
Recruitment of P. subtriangulata is sporadic, and the abundance of cohorts from different years can vary greatly (Greenway 1969, Redfearn 1974, Grant 1994).
Newly recruited clams can occur in high densities (up to 4cm−2) in the intertidal (Greenway 1969, Grant 1994).
Fishery
P. subtriangulata have been harvested in traditional, commercial and recreational fisheries. Significant traditional harvest of P. subtriangulata is suggested by the
large numbers of shells in coastal middens (Cranfield et al. 1994). Commercial landings of P. subtriangulata are substantial and have increased in recent years. The
landings are primarily from northwestern North Island, the area that formerly supported the commercial fishery for toheroa, P. ventricosa. Commercial fishery
landings for P. subtriangulata averaged 53tyr−1 from 1974 to 1978, and 106tyr−1 from 1979 to 1984 (Greenway 1981, Haddon 1988). Recent landings were 83t in
1992, and 131t in 1993 (Fishing Industry Board unpubl. data). Fishers received a port price of US$600t−1 green weight in 1992, and US$480t−1 in 1993.
Tuatua are sold fresh, frozen and as prepared products including “toheroastyle soup” (Greenway 1981) for domestic and export markets. In New Zealand, P.
subtriangulata is marketed in canned products, in jars, and as fresh whole and shucked clam meats for retail sales, catering and restaurants. In 1993, 26t (20% of
landings) of chilled whole clams were
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exported to the USA, and 4t (3% of landings) of frozen clams were exported to Reunion Island and Guam (Fishing Industry Board unpubl. data). The export price for
frozen and chilled whole clams was US$2.65kg−1 in 1993 (Fishing Industry Board unpubl. data).
P. subtriangulata are harvested by hand at low tide. The number of permits issued for commercial take has decreased from 35 in 1981 to approximately six in
1994. There was a limit of three bags (c. 40kg) per fishers per day in 1981. In 1994, the limit was 300kg per fisher per day. Commercial harvest is prohibited in
specified areas near access points for recreational fishers.
Data on recreational harvesting are not available, but landings are likely to be substantial in some areas. Recreational harvesting is unregulated except for a daily bag
limit of 150 clams per person, and closed periods due to toxic algal blooms.
The tuatua Paphies donacina (Fig. 2o, p. 166)
Distribution
The second species of tuatua, P. donacina, is also endemic to New Zealand, occurring primarily along the west coast beaches of North Island, on sandy beaches of
eastern South Island, and on Stewart Island (Fig. 1, p. 164). P. donacina is not known from northeastern North Island and is rare north of Auckland. P. donacina
occurs on dissipative and intermediate beaches, and in some more sheltered locations including harbour mouths. Adult P. donacina are found in the low intertidal and
in the subtidal to depths of 4m, but generally occur shoreward of the primary breaker zone (Cranfield et al. 1994).

The sexes are separate in P. donacina, and occur in a ratio close to 1:1 (Dawson 1954). The eggs of P. donacina are larger than those of P. ventricosa and P.
subtriangulata. Egg diameters of P. donacina are approximately 80μm (Dawson 1954) compared with 60–66μm in the other species.
P. donacina is a suspension feeder with short siphons that are held at the sand surface. P. donacina has slow burrowing rates relative to other beach clams
(McLachlan et al. 1995). Juvenile P. donacina had a BRI of 3 at 15°C. Small clams have been observed surfing in the swash with feet and siphons extended. There is
no evidence for tidal migration by P. donacina.
Predators of P. donacina include southern blackbacked gulls, redbilled gulls, variable oystercatchers and South Island pied oystercatchers. Southern black
backed gulls have been observed diving in the shallow subtidal to extract buried clams of up to 100mm shell length, and opening them by dropping them on to the sand
from a height (J.E.Dugan & D.M. Hubbard unpubl. data). Oystercatchers prey on smaller clams to about 50 mm shell length in the intertidal zone (J.E.Dugan &
D.M.Hubbard unpubl. data).
Richardson et al. (1982) reported that the shells of P. donacina from Wellington west coast beaches were free of hydroids and algal growth, and suggested that
they occur at greater depths in the sand than P. subtriangulata. This may vary geographically as shells of P. donacina from Otago beaches frequently bear hydroid
clumps and algal growth (J.E. Dugan & D.M.Hubbard unpubl. data).
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Population dynamics
High densities of P. donacina recruit to the intertidal where larger clams are rare. Small clams (mean length <10mm) were numerically dominant (>90%) in intertidal
samples from ten Otago beaches (J.E.Dugan & D.M.Hubbard unpubl. data). Samples of P. donacina from the subtidal zone have length frequency distributions that
are primarily composed of a substantial mode of large animals. These modes are probably composed of multiple year classes (Cranfield et al. 1993). The lengths of
clams in these modes were mostly 50–80mm in samples dredged from Wellington west coast beaches, North Island, and 75–105mm in samples from Cloudy Bay,
South Island (Cranfield et al. 1993). P. donacina sampled in the shallow subtidal at Warrington, South Island, had similar length frequency distributions with mean
lengths of 71mm in February 1994, and 78mm in January 1995 (J.E.Dugan & D.M. Hubbard, unpubl. data). P. donacina sampled by Cranfield et al. (1993) reached
larger maximum sizes on South Island beaches (95–109mm shell length at six sites) than on North Island (73–88mm shell length at four sites). The size structures of
subtidal P. donacina populations were stable over time in nine samples from Cloudy Bay and eight samples from the Wellington west coast over 19 months (Cranfield
et al. 1993).
Estimated biomass of subtidal P. donacina from 16 beaches across 11 degrees of latitude was 0–60kgm−1 (Cranfield et al. 1994). Estimated biomass of P.
donacina in the shallow subtidal at Warrington was 1.6kgm−2 (J.E.Dugan & D.M.Hubbard unpubl. data).
Recruitment of P. donacina occurs across the entire intertidal zone (J.E.Dugan & D.M.Hubbard unpubl. data), with substantial variation in recruitment to different
beaches. Estimates of recruit densities (2–12mm shell length) for east coast beaches of South Island (Otago) in 1994 varied from 0 to >3000m−1, and beaches north of
Cape Saunders had much higher densities of youngoftheyear clams (>900m−1) than those to the south (<60m−1) (J. E.Dugan & D.M.Hubbard unpubl. data).
Growth rates and maximum sizes (109mm shell length and 180g WW) for P. donacina are intermediate to those of P. subtriangulata and P. ventricosa (
growth rate index=3.47; Table 2, p. 168). Growth rates of subtidal P. donacina were estimated from a one year markrecapture experiment at Cloudy Bay (Cranfield
et al. 1993). No increase in length was measured for a large proportion of the clams with initial lengths >70mm. It would take 5yr for recruits to reach marketable sizes
(approximately 75mm). Analysis of shell sections indicated that P. donacina reach ages up to 17yr. Cranfield et al. (1993) estimated M of P. donacina at 0.26–
0.32yr−1 for Cloudy Bay.
Fishery
Artisanal, commercial and recreational fisheries for P. donacina are smaller than those for the other beach Paphies species in New Zealand. The commercial fishery
for P. donacina is currently very small, but there is potential for substantial landings if the surfzone clam fishery develops further. One fisher is currently harvesting surf
clams commercially by hydraulic dredge in central New Zealand, but total landings were less than 5tyr−1 from 1990 to 1992 (Cranfield et al. 1994). P. donacina is
one of several species harvested, and composed 20% and 29% of the estimated stock biomass at Cloudy Bay and Clifford Bay, respectively (Cranfield et al. 1994).
Cranfield et al. (1994) estimated maximum constant yield [MCY=0.25 (F0.1) (virgin biomass)] for P. donacina at two sites: 15.5tyr−1 (1.4kgm−1yr−1) from Cloudy
Bay (11km), and 25.8tyr−1 (1.1kgm−1yr−1) from Clifford Bay (23.5km). No data are available on the recreational harvest of P. donacina, but harvesting rates are
probably much lower than for P. subtriangulata. The recreational fishery for P. donacina is unregulated except for a daily bag limit of 150 clams per person, and
closures due to toxic algal blooms.
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Discussion
General
Several clear patterns and contrasts emerge from the foregoing review. Many of the species covered in this review are large (Table 2, p. 168) and found on temperate
dissipative beaches in areas of high primary production, which is often in the form of surf diatom blooms. Species from warmer waters tend to be smaller, occupy a
wider range of beach types and support artisanal fisheries, while large species in temperate areas support or have supported commercial fisheries.
The species covered all come from exposed ocean beaches but can be divided into intertidal and subtidal forms (Table 2), the latter with adults in the surf zones and
the former with adults on the intertidal beach. In both cases juveniles tend to settle at different levels before migrating to the adult zone. A third zonation pattern is
intermediate between these two extremes, with adults in the shallow swash, extending both into the intertidal and out into the surf zone. In the case of one species,
Donax serra, an intertidal distribution occurs in a warm part of its range and subtidal distribution in a colder part. A similar temperature effect is seen in Mesodesma
mactroides, which occurs subtidally in the cold winter months and intertidally in summer. This seems, at least in part, to match the other species patterns: there appears
to be a good relationship between tide level occupied by a clam species and water temperature, those occurring in the warmest waters (Donax denticulatus, D.
striatus, D. faba, D. cuneatus from tropical and Mesodesma mactroides, Donax serra, D. trunculus, D. deltoides, Tivela stultorum from warm temperate areas)
being either intertidal or tidal migrants, whereas those from higher latitudes (Siliqua patula, Mesodesma donacium, Paphies donacina) are mixed or subtidal. The
explanation for this may lie in Donn’s (1990a) hypothesis that the subtidal shift in Donax serra populations in areas of lower water temperature is a consequence of
slower burial rates at these temperatures and thus inability to maintain position in the dynamic swash conditions of the intertidal zone. Burial time increases with size in
bivalves (McLachlan et al. 1995) and large species such as these may be extremely slow at low temperatures. This would make repositioning and burial on the beach
face, an essential process on dynamic beaches, difficult or impossible.
Many of the species covered are large (>50mm) and, unlike many other beach animals, do not undergo tidal migrations. They thus differ from the small, highly mobile
donacids and other species typical of reflective and intermediate beaches, especially in warmer climates. McLachlan et al. (1995) have shown that bivalves from ocean
beaches can be separated on the basis of size, density and shape into forms typical of reflective, intermediate and dissipative beaches. Those from intermediate and
dissipative beaches include large forms with low densities and variable shapes, in contrast to the small, dense, wedgeshaped forms strongly selected to cope with the
turbulent swash conditions of reflective beaches. Donn (1990b) found intertidal populations of D. serra to have higher densities, and smaller, thicker, lessrounded and
more wedgeshaped valves than subtidal populations. Some of the bivalves listed here clearly fall into the category of large species with low densities and subtidal
distributions. The exceptions are probably D. serra on the south coast of South Africa, D. deltoides in Australia, and D. trunculus in Europe, which are intermediate
between the bulky forms from dissipative beaches and the small, dense forms from reflective beaches such as those found in the tropics. This may explain the ability of
these three species to colonize a fairly wide range of intermediate beach types and to maintain intertidal positions.
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The species supporting substantial beach clam fisheries in temperate areas tend to be larger, longerlived and with higher growth rates than many other beach
bivalves (Table 2, p. 168). Many have life spans exceeding 5yr and can attain high biomass values, contributing significantly to biomass on the beaches where they
occur. Most occupy central positions in beach food chains, often constituting the dominant prey of birds, fishes and crabs. But the impacts of clam beds on beach
ecology may extend beyond their role in energy flow. Although competitive interactions have not been demonstrated in exposed sandy beaches, the presence of
substantial numbers and biomass of large filter feeders may be expected to affect other species on the beach. Competitive interactions in sediments are generally not
strong because of the three dimensional space and the effectiveness of predation in keeping numbers below carrying capacity (Peterson 1979). However, de Alava
(1993) has shown in the case of Mesodesma mactroides that competition with Donax hanleyanus may occur since the latter exhibits recruitment fluctuations inversely
correlated with both abundance of juveniles plus adults of Mesodesma mactroides and also the amount of fishing effort focused on the latter (Defeo & de Alava,
1995).
The role of densitydependent processes in structuring exploited sandybeach populations must not be neglected. Defeo (1993a) demonstrated reduced recruitment
under high adult density, presumably due to filtration of larvae on a microscale. The strength of a newly settled cohort could also be inhibited by densitydependent
growth and mortality rates. Depression of growth rates at high density could also have an effect on subsequent recruitment events produced by these cohorts. For
example, cohorts with lower growth rates would not achieve the size at maturity during the spawning season, thus affecting fecundity and possibly leading to recruitment
failure (Defeo et al. 1992b). Conversely, at low adult densities growth rates may be higher and the consequent size at maturity may be achieved during the normal
spawning season.
Population dynamics of species supporting beach fisheries are not particularly well studied and in many cases data on recruitment, stock size, mortality and growth
are incomplete or absent. The two North American clams, Tivela stultorum and Siliqua patula, Donax serra from South Africa, Paphies ventricosa from New
Zealand and Mesodesma mactroides from Uruguay, have been best studied in this regard. Recruitment is probably the least studied aspect of beach clam fisheries and
has been quantified in only a handful of cases. It has been shown to be stock dependent on a macroscale in M. mactroides but may be independent in Tivela
stultorum. Most species have rapid growth to sexual maturity and then slower but nevertheless indeterminate growth and various growth models have been
successfully applied (Table 2, p. 168). Mortality rates vary considerably between species, beaches and years and total mortality can be high, ranging 0.5–3.1yr−1 in
different situations. Fishing mortality makes up a significant proportion of this in some cases and associated incidental mortality (clams not taken but discarded on the
surface to die) can be a major cause of mortalities. Survival probably increases with age in all cases, especially the longlived species.
Human utilization of beach clams extends far back into prehistory on most coasts. Today beach clam harvests seem to be clearly divided into recreational and
commercial fisheries, with a clash of interests between the two in many cases. The scenic nature and sporting potential of beaches renders them ideal for the
development of recreational activities, and recreational fisheries have even replaced commercial fisheries (especially for species in areas with high human density, e.g.
Siliqua patula, Tivela stultorum) as beach recreation becomes more popular and human population increases. Recreational activities on beaches, are not, in general
conducive to coexistence with commercial fisheries, especially for intertidal species in warm climates (Donax serra, D. deltoides) where beach recreation is
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extremely popular. For subtidal species this appears less of a problem since fishing effort is based below the intertidal beach. While all fisheries include some
recreational components, artisanal fisheries predominate in South America and some tropical areas. Commercial fisheries, which in the past were strongly developed in
the case of Siliqua patula, are still important only in the case of Mesodesma donacium in Chile. It appears that the tendency in beach clam fisheries is towards more
recreational/artisanal fisheries and away from large commercial operations, although the former may have commercial interests.
Intertidal species are readily accessible and harvest incurs very low operating costs, thus they are particularly vulnerable to overexploitation and require careful
management. In contrast, subtidal species occupying high energy surf zones are relatively inaccessible and less management is required. In recreational fisheries, limits to
the number of fishers are generally not possible and managers have made recourse to size and bag limits, restrictions on collecting gear and closed seasons and areas.
However, for many of these fisheries research and management are limited or nonexistent. In a few areas restocking or enhancement has been tried as a management
tool but, on the whole, the idea of reseeding beaches has not been tested and in most cases little is known of larval biology. Only on the northwest American coast does
management seem well developed.
Beach clams are used for food and bait. They obtain best prices as food and clams can reach US$30kg−1 in their shells. In areas with restricted access to markets,
limited demand for clam meat, or high occurrence of toxic algal blooms, a significant proportion of the take goes to bait for crabs and fishes in other recreational or
commercial fisheries.
A problem affecting commercial fisheries that target beach clams for eating is the tendency of several species to accumulate phytoplankton toxins produced by
blooms of certain species of phytoplankton. High levels of toxins, such as PSP and domoic acid, are harmful to consumers of bivalves and pose health hazards to
humans and other animals. This limits the potential utilization of many stocks in cooler waters and creates the need for careful monitoring and management. As human
impacts on coastal waters continue, blooms of toxinproducing phytoplankton could affect beach clam fisheries more (e.g. Hallegraeff 1993).
In general it appears that beach clam fisheries are ubiquitous features of open coasts in all latitudes. Despite their extent they are relatively small in comparison with
some commercial clam fisheries. For example the offshore fishery for “surf clams” (Spisula solidissima) and quahogs (Arctica islandica) off the east coast of the USA
had landings of 56000t and a first sale value of US$58 million in 1986 (Murawski & Serchuk 1989). Total beach clam takings for all species covered in this review is
not known but is today certainly below this figure, although in the past landings of Tivela stultorum were comparable.
Future research needs

Based on this review, we suggest that a number of topics deserve special consideration in order to provide a sound scientific basis for management of beach clam
fisheries.
Recruitment processes
Little is known about dispersive abilities of meroplanktonic larval phases of exposed sandy beach macrofauna (Efford 1970) and the mechanisms influencing larval
distribution are still poorly understood. Research should focus on planktonic stages and the role of near shore hydrodynamics in settlement processes. This will
determine the spatial scales at which population dynamics may be considered an open process, i.e. if it is more related to the arrival
Page 220
rates of larvae than to postsettlement processes. Efford’s (1970) hypotheses should be tested to determine whether exposed sandybeach populations could be
considered as selfsustaining, with relative isolation of the rest of the species distribution. The inclusion of physicaloceanographic information related to larval dispersal
would be of importance in this context. In addition, potential mechanisms associated with overcompensatory processes in stockrecruitment relationships, such as
passive filtering of larvae by adults, should be examined by field and laboratory studies.
Densitydependent processes Spatial and temporal variations in growth, mortality and recruitment rates in sandybeach populations may be related in part to density
dependent processes in sandybeach populations (Defeo, 1993a, J.E.Dugan unpubl. data), and hence could affect the clam stock available for exploitation (Defeo et al.
1992b). However, as these processes are relatively unknown for exploited beach clams, future work should emphasize scaledependent experimental manipulations of
resource abundance and population density.
Disturbance associated with harvesting and incidental mortality The natural mortality rates of the unharvested fraction of exploited sandybeach populations may
be directly related to the amount of fishing effort (Schink et al. 1983, Lassuy & Simons 1989, Defeo, 1993a), suggesting that harvesting activities constitute a source of
sediment disturbance and incidental damage particularly in intertidal species. Furthermore, incidental effects (shell damage) of fishing pressure could affect harvested
and unharvested populations in sandy beaches (Defeo & de Alava 1995, Sims pers. comm.). Such shortterm effects need to be further tested through direct
quantification of incidental damage and mortality, as in the Siliqua patula fishery where wastage is reported (Ayres & Simons 1992), and the effects of substratum
perturbation (e.g. mortality of clams immediately after fishing manipulation versus control areas). In clam fisheries where harvesting produces significant incidental
mortality, the reduction of minimum size limits may reduce losses of small clams (see Lassuy & Simons 1989). Such regulatory measures could be experimentally
evaluated in vulnerable clam stocks. In order to evaluate the longterm effects of harvesting on the population dynamics of the target species and other species within
the community, human exclusion experiments (see Defeo, 1993a) could be used.
Modelling and management Management must account for temporal and geographic variation in life history and population dynamics characteristic of fished stocks,
e.g. Mesodesma donacium of Chile, Donax serra of South Africa. Since the allocation of fishing effort closely follows spatiotemporal variations in resource
abundance, fishing effort and catches should also be obtained discriminately by homogeneous areas (see Defeo et al. 1993 and references therein). The spatially
discrete analysis of sandy beach stocks, the surrounding environment, and the fishery, will be a useful tool for (a) the assessment of spatial dynamics of catch and fishing
effort, (b) the detection of changes in length or age composition of the catch, and (c) monitoring changes in stock abundance and its composition by size and/or age, as
well as in other population dynamics parameters and environmental variables. From the above, an integrated approach could be built to develop a comprehensive
management scheme of the fishery (Defeo et al. 1993). Since aquaculture has not been feasible in most of the species reviewed, and considering their increasing
demand, we recommend an experimental approach to management through rotation of fishing areas, creation of spawning stock refuges and restocking through direct
seeding of juveniles or adults (Castilla 1988, Defeo 1993b, Dugan & Davis 1993). A spatial management scheme could complement these
Page 221
strategies by accounting for spatial variation in the magnitude of recruitment, and thus protecting those areas with high probabilities of successful recolonization (Caddy
1989).
Future prospects
Although beach clams do not provide very large commercial catches they are ideally suited to recreational exploitation because of the popularity of beaches as
recreational areas. Ready accessibility, large size and slow growth of several temperate species does, however, necessitate careful management to prevent
unsustainable exploitation. The same may hold for tropical species, especially in areas where large numbers of poor people depend on these for subsistence.
Unfortunately, beach clam stocks have often received limited attention by authorities and fisheries managers, as evidenced by the paucity of information reviewed. It is
important that these stocks be seen as a valuable resource even where they do not support commercial exploitation. The value of recreational fisheries lies not only in
the value of the meat but also in the recreational enjoyment of the whole experience. Thus the true value of beach clam stocks may include a small food component and
a large recreational activity component. This must be recognized and managed so that beach clammers can be assured of a quality recreational experience even if their
take is modest.
Overexploitation by recreational and commercial interests is not the only threat facing beach clam fisheries. Any threat to beaches in general also threatens the
stocks. Offroad vehicles, pollution, heavy recreational pressure, coastal engineering structures, beach erosion or replenishment programmes and coastal development
all pose problems. With increasing global population and sea level rise, both directly impacting on ocean beaches, these stocks must be assumed vulnerable and the
formulation of dynamic management strategies is essential.
Acknowledgements
JED was responsible for the sections on T. stultorum and S. patula, PEP and ADA for T. mactroides, D. denticulatus, and D. striatus, OD for the section on M.
mactroides, EJ for that on M. donacium, ADA for D. trunculus, D. cuneatus and D. faba, AM for D. serra and D. deltoides, and DH for the Paphies species. We
thank the following who provided information and assistance with the section on S. patula: D.Ayres, D.Simons and A.Rammer (Washington State Dept. Fish and
Wildlife, USA) and N.Bourne (Fisheries and Oceans, Pacific Biological Station, Canada). D.del Pietro (Università di Trieste, Italy) and C.Salas (Universidad de
Málaga, Spain, provided information on D. trunculus and N.Hacking assisted in obtaining information on D. deltoides. H.J.Cranfield (MAF), G.Bremner (MAF
Fisheries South), G.Mason (University of Otago), G.Stace (Auckland Museum) and R.Creese (University of Auckland) assisted with information for the New Zealand
section. We are grateful to Doug Simons, Washington State Department of Fisheries and Wildlife, USA, and to Simon Grosset, photojournalist, for permission to
reproduce the photographs in Figures 7 and 11, respectively.
Page 222
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BIOLOGY OF THE GASTROPOD FAMILY LITTORINIDAE.

I. EVOLUTIONARY ASPECTS
CHRISTOPHER D.McQUAID
Department of Zoology and Entomology, Rhodes University,
Grahamstown 6140, South Africa

Abstract Gastropods of the family Littorinidae are abundant and ecologically important animals in shallow marine systems throughout the world.
Ecological studies on this family have focused on geographical areas where the taxonomy is clear at species level. In some areas, notably the North
Atlantic, this happy state does not exist and there has been extensive taxonomic confusion. As a result, considerable effort has gone into the study of
the evolutionary biology of the littorinids over the last 20 years. Many of these studies have been aimed at understanding the taxonomic status of
species. I have not addressed these. Others have used the group as a tool to consider evolutionary questions. These concern the maintenance of
polymorphism in shell colour or shape, and especially the application of life history theory to mode of larval development and the influence of
development type on dispersal and population heterozygosity. Overall the littorinids conform poorly or not at all to theoretical predictions.
Colour polymorphisms seem to be maintained by selective forces based on predation (although the ecological literature suggests predation is not
important at the population level) and heat uptake. The control of variation in shell shape is more difficult to explain and is influenced by the type of
development. Direct developing species lack a planktonic larva, which is assumed to result in low gene flow. In examples of these species, shape
appears to be largely genetically determined. For species with highly dispersive larvae, phenotypic responses to local factors, including growth rates,
are important. But this generalization is not clearcut, the evidence is conflicting and important counterexamples exist. The most direct evidence comes
from breeding experiments. These are rare and, in at least two cases, provide examples that do not conform to this pattern.
Dispersal and gene flow are expected to correlate with mode of larval development. In fact, we have little direct information on events between
spawning and settlement and again this link is not clear. Direct developing species have been shown to raft or drift over long distances as adults.
Species with longlived planktonic larvae are expected to be highly dispersive, but correlations between settlement and adult density can be interpreted
as indicating limited dispersal. A number of other observations on actual dispersal and geographic range of littorinids with different larval types do not
fit the theory.
Direct developing species can show striking examples of founder effects. Despite this, there are only imperfect correlations between larval type and
both within and amongpopulation genetic heterozygosity, implying that gene flow is not tightly bound to larval type. This may be partly explained by
the lack of correlation between larval type and dispersal. We may also be misled by assuming that juvenile recruitment is not genotype dependent. But
most important is the fact that, while development mode is fixed for any species, heterozygosity is influenced by many factors. The absence of a clear
correlation suggests that larval type and/or dispersal is only one of many factors that influence heterozygosity and does not have an overriding effect.
There is a poor fit between predicted and actual reproductive strategies used by littorinids. One of the most useful ideas to emerge recently is the
need to recognize phylogenetic constraints on mode of reproduction (and many other facets of biology for that matter). Evolutionary explanations can
supersede a welter of imperfect and superfluous ecological explanations for the use of different strategies by different species. We require adaptive
explanations only when a species shows a derived condition different from the primitive condition for the taxon to which it belongs. Thus ecological
explanations are appropriate only in comparisons between species of similar ancestry or between populations of the same species exhibiting
morphological or genetic differences.
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Introduction
In the preface to the Proceedings of the Third International Symposium on Littorinid Biology, the editors observed that members of the Littorinidae are probably
among the most thoroughly studied of all marine gastropods. Certainly the literature on these animals is formidable. My original intention was to review the literature on
the biology of the family, a task that quickly proved beyond me. In the end I have focused on two broad areas of research. The first (this paper) deals with evolutionary
aspects. Here I do not address taxonomic issues, but review information on gene flow, polymorphism, and life histories. Central to this is the supposed link between
mode of larval development and dispersal. The second (McQuaid 1996) covers the role of littorinids in shallow marine ecosystems. This does not include autecological
or ecophysiological issues, but rather interactions within and among species including algae, other grazers, predators, etc. At the end of each paper I attempt to define
the primary outstanding problems in these two fields and offer directions for the most profitable ways of addressing them.
It is noticeable that different species, or higher taxa, attract different degrees of attention partly because of their obvious (or apparent) ecological importance and
partly because more biologists live in some parts of the world than others. Different taxa are also associated with different types of research. The North Atlantic has a
relatively young littorinid fauna, and ecological research on many species or species complexes has been crippled by taxonomic confusion. These animals attract
attention from taxonomists and from evolutionary biologists. The high degree of variation and taxonomic confusion within the genus Littoraria has focused attention on
mechanisms responsible for maintaining colour polymorphisms. Littorina littorea has precipitated a huge amount of ecological research in North America because of
its central place in the ecology of rocky shores in the northeastern USA. It is safe to say that different types of research predominate in different geographic areas. This
largely reflects where different species occur. Perhaps, too, biologists in different parts of the world have different fields of interest.
Taxonomy and evolution

The taxonomy of the Littorinidae has been problematic at both specific and supraspecific levels. For nontaxonomists the latter can seem esoteric and ecologically
irrelevant, but a phylogenetically based taxonomy can provide solutions to intractable ecological problems. An excellent example is given by studies of “life history
strategies” that are discussed below. The most recent analysis of the taxonomy of the family is given by Reid (1989) who uses a cladistic approach. His cladogram is
based on 53 characters including features of the shell, operculum and radula that formed the basis of previous classifications (e.g. Rose water 1970, 1981), as well as
the reproductive system and larval development. The results differ somewhat from previous schemes, particularly in the number of subfamilies and the arrangement of
genera within them (cf. Tables 1 and 2 in Reid 1989). The cladogram indicates that the family Littorinidae is monophyletic, with three subfamilies. The subfamily
Laevilitorininae may be paraphyletic or monophyletic. It comprises a number of less well known genera limited to the Antarctic and southern Australia or New Zealand.
The other two subfamilies are monophyletic and include a number of conspicuous or ecologically important species. The Lacuninae includes Lacuna, an important
northern
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hemisphere grazer and Bembicium, which is common on Australian rocky shores. The third subfamily, the Littorininae, contains the best known genera—the
mangrovedwelling Littoraria and two common rocky shore genera, Nodilittorina (the most species rich genus in the family) and Littorina itself.
Reid (1989) suggests a southern temperate origin for the Lacuninae and Laevilitorininae, with Lacuna spreading to its present range in the north temperate via
relatively deep, cool water along the west coast of the Americas and more recently into the Atlantic. The Littorininae, however, are tropical or temperate. Littoraria is
pantropical and, while there are two clear centres of high endemicity and richness in northwestern Australia, and the Malay peninsula and Borneo area (Reid 1986), the
geographical origins of the genus are unclear (Reid 1989). Littorina may have originated in the Tethys Sea, spreading west to the Pacific and later reinvading the
Atlantic via the Bering Straits (Reid 1990a).
The need for a sound taxonomic basis at the species level is more obvious. It is impossible to understand ecological or adaptive responses unless one can
differentiate between groups of species and single, but highly polymorphic, species. For example, Reid (1986) divides Littorina scabra into 17 species of Littoraria.
Consequently, apparently adaptive responses of one species may now be understood as evolutionary radiation of a number of species. The difficulties caused by an
inaccurate taxonomy can be serious and are classically demonstrated by the case of European Littorina spp. This problem involved the taxonomic status of a
bewildering array of real or apparent ecomorphs and synonyms, especially within the Littorina saxatilis complex, and is reviewed by Smith (1982) and Raffaelli
(1982, 1990). Until the work of Sacchi & Rastelli (1966), four European species were recognized. After several permutations, it seems to be generally accepted at
present that L. littoralis includes L. obtusata and L. mariae, while L. saxatilis has been split into four species: L. saxatilis, L. arcana, L. nigrolineata, and L.
neglecta. The issue is not yet resolved as the specific status of L. neglecta has been challenged (B.Johannesson & Johannesson 1990, K.Johannesson & Johannesson
1990, Reid 1993, but see Grahame et al. 1992). Consequently, we can make nothing of earlier studies that did not distinguish between these various species. For
example, the findings of Beardmore & Morris (1978) and Wilkins et al. (1978) that L. saxatilis and L. littoralis exhibit greater genetic heterozygosity than L. littorea
may mean that they included several species under a single name.
Defining species
The genetics of some littorinids have been studied intensely over the last 15 to 20 years not only for taxonomic reasons, but also because of interest in life history
strategies. Johannesson & Sundberg (1992) consider the relationship between polymorphism and incipient speciation in L. saxatilis. Essentially, the question of specific
status concerns defining gene flow between populations. Analysis of morphological differences does this very indirectly and is confounded by phenotypic responses.
Ultimately breeding experiments provide the final answer to problems of specific status, but these can be difficult and are not widely used. Less direct measures, using
genetic techniques, are commonly used to resolve taxonomic questions. These can be problematic for two reasons. First, both microgeographic and temporal effects on
phenotypic variation make the method of sample collection critical (B.Johannesson & Johannesson 1990). This, or variations in laboratory technique, may explain the
fact that different authors can obtain radically different results. Examples are given by genetic distances
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measured between L. obtusata and L. mariae. These range between 0.087 (Morris 1979, in Ward 1990) and 0.501 (Moyse et al. 1982). A more worrying case is
comparison of heterozygosity in European and North American populations of L. littorea. American populations have been found to be more, less, and equally
heterozygous to European populations by various authors (Vuilleumier & Matteo 1972, Berger 1977, Morris 1979, in Ward 1990). Clearly these authors were
measuring different things, but this serves to show that genetic techniques are not infallible. The second problem is that, even with an accurate estimate of genetic
distance or identity between two populations, there is still the decision about how dissimilar two things must be in order to be different.
For example, L. arcana has a different reproductive mode from L. saxatilis (Fretter & Graham 1994), sympatric populations show different distributions of allele
frequencies (Ward & Janson 1985), and the two are clearly different species. Yet Janson (1985a), I believe rightly, accepts populations of L. saxatilis from Venice as
the same species as nonMediterranean populations because of a high genetic identity. In fact this identity is lower than that of L. arcana to North European L.
saxatilis (Ward & Warwick 1980, Knight et al. 1987, Knight & Ward 1991). We can explain this in terms of founder effects, but it highlights a classic dilemma for the
biological species concept: how to deal with geographically separated populations. Genetic differences between sympatric populations tell us that there is little or no
interbreeding. Differences between allopatric populations can be due to purely geographic effects and do not tell us if interbreeding is possible.
Despite these problems a number of authors have successfully used electrophoretic techniques to clarify species relationships in terms of genetic differentiation (e.g.
Ward & Warwick 1980, Mastro et al. 1982, Janson 1985b, Janson & Ward 1985, Knight & Ward 1991, Boulding et al. 1993) and the use of allozyme studies to
elucidate taxonomic difficulties is reviewed by Ward (1990).
The principle of electrophoresis is that proteins have characteristic rates of migration in an electric field. As different alleles produce different proteins their presence
can be deduced by measuring the mobility of protein extracts. Even this is not entirely direct and involves the inference of genetic variability from enzyme variability.
Such studies should ultimately be supported by breeding experiments, which are rare so far (e.g. Warwick 1983, Ward et al. 1986, Boulding & Hay 1993, Boulding et
al. 1993, LanganCranford & Pearse 1995). Accepting this proviso, data on allele frequencies allow comparisons of implied gene flow between morphs and between
geographically separated populations, but Beaumont (1991) notes two potential pitfalls. First, in the case of nonsympatric populations, different selective pressures
could cause different genotypedependent mortality, producing different allele frequencies. Janson & Ward (1984) found, with the exception of one locus, little
evidence for the importance of selection in maintaining allozyme variability, despite selection for shell shape according to the degree of wave action. Johannesson &
Johannesson (1989), however, give evidence for selection of particular alleles of one enzyme locus in mid and uppershore populations of L. saxatilis. Similarly,
Newkirk & Doyle (1975, 1979) found a correlation (in two estuaries) between allele frequencies and distance from the estuary mouth. The second problem is still
more intractable; there may be real genetic differences between populations, but at different loci from those studied.
Encouragingly, allozyme studies tend to corroborate the cladistic analyses. The results of Janson (1985b), Ward (1990) and Backeljau & Warmoes (1992) all agree
with the predictions of the phylogeny of Reid (1990a). A more recently developed technique is randomly amplified polymorphic DNA (RAPD) analysis that has been
used to separate L. saxatilis from L. arcana more successfully than has been possible using enzyme polymorphisms (Crossland
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et al. 1993). This technique is likely to be useful in both taxonomic and population studies in the future.
Morphological variability
One attribute of some littorinids that has made them an attractive subject for research is the morphological variability of their shell characters, particularly shell shape
and the existence in many species of different colour morphs. This can occur both within and between populations; I shall use the term polymorphism for both. There
are difficulties in the search for the cause of morphological patterns. Bandel & Kadolsky (1982) point out that there are many selective processes operating on
morphology, both biological (predation, finding mates, etc.) and physical (heat stress, desiccation, wave action, etc.). Also, each of these problems can be addressed in
several different ways, so that morphology reflects a complex pattern of adaptation. Simple explanations may be valid in narrow ranges but cannot often be generalized.
Perhaps a more optimistic view is that, despite many exceptions, there are some generalities that appear to have some validity. For example, Vermeij (1973) noted a
general upshore increase in shell ornamentation and decrease in shell aperture size in tropical littorines and correlated these with increasing heat and desiccation stress.
There are many exceptions to this pattern and yet it is often applicable (e.g. Bandel & Kadolsky 1982). A survey of research on this topic brings out the importance of
both genotypic and phenotypic effects. In the case of shell shape, their relative importance may depend on the mode of development of the species.
Shell colour
Shell pigments of archaeogastropods are proteinbound and probably formed as digestive residues or aborted synthetic pathways (Cole 1975). Consequently, both
colour and patterning can be determined by the availability of food (e.g. Ino 1949, Leighton & Boolootian 1963, Creese & Underwood 1976, Underwood & Creese
1976). The pigment system of meso and neogastropods is fundamentally different (Comfort 1951). Their shell colour may not be entirely independent of diet (e.g.
Cole 1975), but colour polymorphism has been shown through breeding experiments to have a genetic basis in L. mariae (Reimchen 1979), L. saxatilis (as L. rudis)
and L. arcana (Atkinson & Warwick 1983). Consequently, shell coloration in littorinids has been used to investigate evolutionary processes.
The case of polymorphism in the land snail Cepaea has been offered as a caveat by Jones et al. (1977). They review a very extensive literature and show that at
least eight evolutionary forces operate on shell colour polymorphism in Cepaea. In most populations they operate simultaneously, their relative importance depending
on the locality, and it is rare for any one factor to explain fully observed frequencies of colour patterns. Jones et al. (1977) conclude that each population requires a
complex and possibly unique solution to the problem and that it is meaningless to seek a single answer. The selective processes implicated in maintaining colour
polymorphism in littorinids are predation and heating.
When migration can be excluded and withinpopulation mating is random, differences in genotype frequency between populations are considered to be due to
selection (Milkman 1978). However, this assumes random settlement of larvae, which is not always the case and
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may even be exceptional. Selective mortality of larvae and hydrographic isolation, which minimizes larval exchange between populations, may maintain genetic
differences between stocks of the mussel Mytilus edulis (Levinton & Lassen 1978). Furthermore, mating may not be random in populations of Littorina (Janson
1982a, Johannesson et al. 1993), but there is still substantial evidence for the effects of selection. Kalabushkin (1976) compared morph frequencies of contemporary
and Holocene L. squalida at Sakhalin Island and found no significant changes over 2000 generations. He suggests that differences between subpopulations are
maintained by selection processes occurring shortly after settlement. Sacchi (1979) correlate the frequencies of various morphs of L. saxatilis with the degree of
exposure and concluded that the main colour morphs can be accepted as ecological indicators. A number of other authors have correlated proportions of colour
morphs with the background colour of the substratum. As birds and crabs are the most significant predators of many littorinids, including Littorina spp. (Pettitt 1975),
differences among habitats have been attributed to crypsis and the intensity of predation by visual predators (e.g. Heller 1975, Smith 1976, Naylor & Begon 1982,
Atkinson & Warwick 1983, Cook et al. 1985, Janson 1985c, Hughes & Mather 1986, Byers 1990).
Frequencydependent predation has been suggested as a mechanism for maintaining polymorphism within species (Clarke et al. 1978, Endler 1978, Cook 1983,
1990), but morphdependent differences in rates of predation have rarely been demonstrated. This is understandable as Endler (1978) lists seven factors that can
determine the pattern producing the best camouflage. These include attributes of the predator (visual acuity, colour vision, predation intensity, etc.) as well as of the
environment (background heterogeneity, etc.) while particular morphs may be selectively neutral if any random selection of the background pattern is cryptic. Although
he did not measure predation per se, Reid (1987) showed frequencydependent disappearance of colour morphs of Littoraria filosa from mangrove trees. Likewise,
Hughes & Mather (1986), working on Littoraria spp. (as Littorina spp.) living on mangrove trees, showed higher survival of yellow morphs unless bird and fish
predators were excluded by caging. Both of these papers, and that of Atkinson & Warwick (1983), are taken as indirect evidence of apostatic selection, that is
selection which favours the less common morphs as predators specialize in capturing the more common prey types. Mercurio et al. (1985) have shown enhanced
mortality of limpets mismatched with different colours of substratum, but the most convincing evidence for the importance of colour in the mediation of predation,
indeed the only work measuring predation directly, is that of Reimchen (1979, 1989) who measured predation of juvenile Littorina mariae by fish (Blennius pholis) in
the laboratory. In the first of these papers Reimchen examined predation of yellow (citrina) and brown (dark reticulate) morphs on natural substrata of different colours.
Juvenile Littorina mariae occur on the kelp Fucus serratus and on brown lamina and stems the yellow citrina was preferred. However, when light was transmitted
through the lamina (i.e. when viewed from below, the normal perspective for blennies) the lamina appears yellow and predator preferences are reversed. In the second
study, relative predation of a white spiralled morph again depended on the background and decreased amongst spirorbid polychaetes. Reimchen (1989) suggests that
this is a case of mimicry.
A second possible source of morphdependent selection is heat uptake and tolerance. Differential heating and habitat use by colour morphs has been observed for
several species, including Littorina saxatilis (Sergievsky 1992). Cook (1986) used correlative data to suggest that different morphs of Littoraria pallescens actively
select different habitats at higher air temperatures. Under a given regime of radiation the final body temperature was lower for paler morphs (Cook & Freeman 1986).
Nodilittorina africana also shows morphdependent
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differences in behaviour in response to increasing temperature. In both cases, differences in behaviour between morphs may be due to different body temperatures at
given intensities of radiation, but they do produce different microhabitat selection in both the field and in the laboratory (Cook 1986, McQuaid & Scherman 1988).
Reid (1987) measured body temperatures of different colour morphs of Littoraria filosa in the field and found slight differences, but all temperatures were well below
probable lethal limits and he suggests that climatic selection may be insignificant. Cook & Garbett (1992) found no link between survival and colour of marked L.
pallescens in the field, but colourdependent mortality linked to heat stress has been found in the laboratory for Nodilittorina africana (McQuaid & Scherman 1988)
and in the field for Nucella lapillus (Etter 1988).
Finally, indirect evidence for the importance of selective processes is given by sympatric species showing apparent convergent evolution of morphological types
characterized by colour and/or shape when living in similar habitats. This has been suggested for Nodilittorina angustior and N. lineolata (Janson 1985b). It also
occurs in “barnacle morphs” of Littorina neglecta, L. nigrolineata and L. arcana (Reid 1993).
Shell shape
The predominance of particular shell shapes, including shell thickness, in different populations has been ascribed to genetic differences, phenotypic differences, growth
rates, and differences in population size composition. The basis for differences in shell shape may depend on the mode of reproduction of the species. The balance of
evidence is that shell shape is largely determined genetically in L. saxatilis, which is a direct developer, and as a nongenotypically based phenotypic response in L.
littorea, which has pelagic larvae. But there are counter examples for species with either mode of reproduction. There is also a suggestion that shell shape may vary as
a consequence of character displacement in populations of L. arcana which cooccur with L. saxatilis, although the selective pressures that may be involved are
unknown (Grahame & Mill 1989).
Direct developing species

Species with nondispersive larvae are assumed to experience reduced gene flow (see below) and habitatspecific morphological or allozyme differentiation in direct
developers has been taken as evidence for differences in selective regimes on scales as small as metres (Tatarenkov 1995, Johannesson et al. 1993, Tatarenkov &
Johannesson 1994). These two facts suggest that consistent morphological differences between habitats, which exist on a wide range of scales, represent genetically
based ecomorphs maintained by selection.
Littorina saxatilis L. saxatilis shows convergence of shell shape in geographically separate populations living in similar habitats and this can be attributed to
convergence of selective pressures. This occurs in the “tenebrosa” phenotype (Janson & Ward 1985, Ward et al. 1986) that is found in brackish water populations as
far apart as Venice (Janson 1985a) and saltmarshes in New England (B.Johannesson & Johannesson 1990). A more intensively studied case is the link between shell
form and wave exposure. Shell shape and thickness generally covary and, in the case of L. saxatilis, this is perceived as
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being in response to similar selective pressures associated, directly or indirectly, with the degree of wave action. The argument is given with numerous references by
Raffaelli (1982).
Usually populations on exposed shores have a relatively larger aperture and thinner shell. Animals with a larger foot can attach to the substratum more strongly (e.g.
Hylleberg & Christensen 1977, McQuaid 1981, Trussell et al. 1993) and a large aperture is thought to be linked to foot size and the ability to withstand wave action
(Raffaelli 1982, Grahame & Mill 1986). On sheltered shores, shells tend to be thicker and have smaller apertures (Raffaelli 1982 and references therein). The strength
of a shell depends directly on it thickness, and possibly microstructure, rather than ridging (Brandwood 1985, Cook & Kenyon 1993, Lowell et al. 1994), and
selection on sheltered shores is thought to favour resistance to either crab predation or, on boulder shores, crushing by rocks (see Raffaelli 1982). This is supported by
the fact that, in the laboratory, crabs (Carcinus maenas) are less successful at attacking thickshelled Littorina saxatilis with small apertures and actively select thin
shelled snails from cliffs rather than thickshelled animals from boulder fields (Johannesson 1986). Boulding & van Alstyne (1993) moved Littorina sp., which has a
thin shell and large aperture, from exposed to sheltered sites. Survival was low and recovered marked shells frequently showed signs of crab predation. The situation is
complicated by cooccurring L. saxatilis and L. nigrolineata showing opposite responses on adjacent boulder shores and cliffs (Smith 1981, Nay lor & Begon
1982). Some authors also disagree on the existence of sexbased differences in some species or size dependent effects in L. saxatilis (Raffaelli 1979, 1982, Van
Marion 1981).
Confusion on size effects may be due to inadequate definitions of size or shape. Shell shape in L. saxatilis is at least partially under genetic control (Newkirk &
Doyle 1975, Janson 1982a,b) and exposed and sheltered morphs have been described that are genetically based ecotypes (Janson 1982a, Janson & Sundberg 1983)
and which also show differences in their tolerance of salinity (Sundell 1985). Sundberg (1988), however, criticizes the arguments given above on shell shape as
correlations have been between shell shape and exposure, rather than the supposed selective agents of predation or risk of wave removal. He also argues that sizefree
comparisons between populations using ratios or regressions are misleading as they are mathematically inadequate. Sundberg uses multivariate techniques to show that
differences between populations subject to contrasting regimes of exposure are mainly due to a general size factor, animals being smaller on wave exposed shores.
Trussell et al. (1993) found the same for L. obtusata. Janson (1982b) used translocation experiments to demonstrate genetic differences in growth rates between
populations, but Sundberg suggests that body size itself may also be selected for and cites Janson’s (1983) evidence of higher mortality for large animals of both
morphs on an exposed shore.
Nucella Studies on this thaiidid whelk modify our conclusions on the genetic control of shell shape. Vermeij (1982a,b) compared the morphology of conspecific
populations in time rather than space, with essentially the same results. Unlike the planktonic developer Littorina littorea (see below), North American Nucella
lapillus have adapted phenotypically since the arrival of the exotic predatory crab Carcinus maenas through an increase in the thickness of the shell lip over the last
100 or so years. Subsequently, Seeley (1986) reported the same response, but more marked, in L. obtusata (another nonplanktonic developer) and demonstrated
preferential survival of thickshelled individuals in areas of high crab density. Vermeij
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(1982b) suggests that locally adaptive phenotypic responses are possible for Nucella lapillus because of reduced gene flow. Seeley also proposed a genotypic
explanation, attributing changes in shell shape to intense selection, but there is an alternative explanation.
Phenotypes are produced by the interaction of genotype and environment and gene flow need not enter into the issue except in its, as yet, unrevealed influence on the
capacity of organisms to respond phenotypically. Although shell morphology of Thais emarginata is under genetic control (Palmer 1982), the phenotypic expression
of the genes controlling shell sculpture can be modified by environmental conditions, such as food availability (Palmer 1985). This finding is reinforced in papers by
Appleton & Palmer (1988) and Palmer (1990). They showed that growth rate and shell form of Nucella spp. were strongly influenced by effluents from predatory
crabs or damaged conspecifics, even when held in separate aquaria. Growth was affected because foraging behaviour was modified by exposure to effluents. Among
starved snails exposed to effluents from feeding crabs, the response was greater if crabs were fed on a diet of Nucella rather than one of fish. An important point here
is that Nucella showed the same response to the threat of predation even if it was not feeding. This suggests that these morphological changes were not due solely to
differences in growth rates. N.lamellosa and N.lapillus show the same pattern of shell thickness in different habitats as littorinids. Animals from sheltered areas (where
there are many crabs) have thicker shells than those from exposed areas (few crabs). In N.lamellosa, thickshelled animals initially showed a stronger response to
effluents than thinshelled individuals. This suggests greater phenotypic plasticity in thickshelled populations that are exposed to crabs in the field, but it could also be
that there is more calcium available for relocation in thick shells. In N. lapillus, however, the reverse was true; thinshelled animals from exposed shores showed a
stronger response. The possibility of such a marked ecophenotypic response reduces the need for explanations of modifications of shell shape based on selection
processes.
Littorina sp. Recent studies by Boulding and coworkers have made an experimental attempt to unravel genotypic and environmental effects on shell form, using
Littorina sp., a directdeveloping northeastern Pacific species related to L. sitkana. Growth rate was found to affect shell form. Boulding et al. (1993) found that
differences in shell shape in split broods of Littorina sp. reared at high and low growth rates were greater than among Littorina sp., L. kurila and L. sitkana. An
important finding by Boulding & Hay (1993) was that positive and negative genetic correlations between features placed constraints on short term responses to
selection. Negative correlations could preclude certain combinations (and thus shell shapes), while additive interactions could produce nonadaptive combinations. In
their study a negative correlation between shell height and weight would promote simultaneous selection for low spired, heavy shells. Both features would increase
resistance to crab predation.
Planktonic developing species

Littorina littorea In the case of L. littorea, which has pelagic larvae, genetic variability between populations is reduced (Janson 1987a, Johannesson 1992) and
differences in shell shape are phenotypic, reflecting differences in growth rates. These in turn vary between
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dense and sparse populations (Kemp & Bertness 1984) and possibly between warm and coolwater populations (Dudley 1980, Vermeij 1982a). Calcification is also
reduced for coolwater gastropods (Vermeij 1978) and this may influence shell shape.
Unlike the poorly dispersed whelk Nucella, Littorina littorea does not show localized adaptation to the presence of predators. Shell thickness does not vary with
degree of wave exposure (Currey & Hughes 1982) and this is reflected in the ease with which crabs can break open snails from either exposed or sheltered shores
(Ash 1989). Similarly Vermeij (1982a,b) found that, again unlike Nucella, North American populations of Littorina littorea have shown no change in shell thickness
since the arrival of Carcinus maenas. He suggests that high gene flow between populations prevents locally adaptive responses in Littorina littorea.
Nodilittorina hawaiiensis In contrast to Littorina littorea, another species with planktotrophic larvae provides strong evidence for genetic control of shell
morphology. Selection for particular genotypes under different conditions of wave action has been shown by Struhsaker (1968) in breeding experiments on Hawaiian
Nodilittorina hawaiiensis (studied as Littorina picta). Phenotypes from different wave regimes breed true, at least to the F1 generation. There is convergence with
European L. saxatilis as waveexposed populations tend to be smaller with lower growth rates and smoother shells than populations from sheltered shores.
Populations from intermediate conditions show intermediate characters and Struhsaker considers this an example of an adaptive polymorphism maintained by
differential selection and nonrandom copulation within two major habitats.
In summary there is evidence for both genetic and environmental control of shell morphology among both poorly and well dispersed species. It seems likely that both
factors are important in all cases, with greater emphasis on genetic control among poor dispersers and on phenotypic response to environmental conditions among well
dispersed species. Breeding experiments address the question most directly and give the more controversial results. Growth rates affect shell shape of direct developing
Littorina sp. (Boulding et al. 1993) while wave actiondependent morphs of the planktonic developer Nodilittorina hawaiiensis breed true (Struhsaker 1968). An
experiment on the type of response examined by Appelton & Palmer (1988), but using a planktonic and a direct developer instead of thick and thinshelled specimens
of the same species would be valuable.
Larval development
Reid (1989) considers the production of benthic egg cases with planktotrophic larvae to be ancestral to the family as a whole while nonplanktotrophic development is
derived. The more primitive subfamilies, the Lacuninae and Laevilitorininae, produce benthic egg cases that release either planktotrophic or nonplanktotrophic larvae.
The theoretical interest in littorinid life histories lies in the subfamily Littorininae, which is characterized by the presence of a capsule gland and the production, at least
primitively, of pelagic egg capsules. Some members of the genus Littorina have evolved ovoviviparity and the genus as a whole exhibits a wide range of developmental
types.
Studies on the reproduction of littorinids have been used to generate or test predictions on three main topics:
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1. Heterozygosity within and among populations

2. Dispersal
3. Life history theory
Broadly, littorinids conform only to theoretical predictions in the first of these topics and even this is tenuous.
Heterozygosity
The genetic diversity of a species results from the balance between factors promoting local differentiation (mutation, genetic drift, local selection) and those promoting
homogeneity (gene flow). In this section I shall consider mainly genetic drift and selection on one hand and gene flow on the other. Genetic diversity has two aspects:
allelic diversity (the number of alleles at a given locus) and heterozygosity (calculated as ; where x i=frequency of allele i). In most of the studies cited here
heterozygosity alone is considered.
Theory
There is considerable controversy about the meaning of protein variability of organisms. Selectionist theory argues that high diversity is maintained by selection. The
assumption is that nucleotide substitutions are fixed because they are advantageous (e.g. Clarke 1970, Lewontin 1974). By this argument, heterozygotes may be “fitter”
than either homozygote (Clarke 1979). However, rates of evolution calculated from rates of nucleotide substitution produce unrealistically high values. This suggests
that many mutations must be selectively neutral (Kimura 1968). Neutral theory assumes that most nucleotide substitutions are selectively equivalent to those they
replace. Protein diversity is therefore attributed to undirected mutation and genetic drift. This theory predicts that heterozygosity is a function of the rate of mutation and
population size. Wilson et al. (1977) suggest that most of the proteins measured using electrophoresis are selectively neutral, but that selection operates on regulatory
rather than structural genes.
These theories have also been extended to include enzyme polymorphism (e.g. Nei & Tateno 1975, Nei et al. 1976) and an interesting modification of neutral theory
by Ohta (1974) suggests that the rate of base substitution is not independent of population size, but is higher in small populations. Thus nucleotide substitution is
concentrated at the time of speciation when small populations create bottlenecks. This could explain some of the anomalies of withinpopulation heterozygosity
observed in comparisons of direct and pelagic developers (see below).
Some of these theories have been applied to littorinids. In order to analyze electrophoretic data, allozyme heterozygosity of littorinid populations is frequently tested
for HardyWeinberg equilibrium. The HardyWeinberg rule predicts that, with random gamete association, genotype frequencies in one generation will follow a
multinomial distribution, where allele frequencies form the distribution parameters. Lessios (1992) outlines serious problems with the interpretation of such studies.
HardyWeinberg equilibrium is only expected under a set of preconditions that mainly concern random combinations of gametes (random mating, infinitely large
populations), absence of selection (genotypes have equal selectivity) and no interference with gene ratios through mutation or migration. Natural populations do not
conform to all these conditions, so the object of comparison of observed and predicted frequencies is to check for violation of the preconditions.
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Ward (1990) summarizes such tests for littorinid populations and finds that, on the whole, they conform to HardyWeinberg expectations. The HardyWeinberg
equilibrium forms the null hypothesis tested by such analyses. Failure to detect a significant difference between observed and expected frequencies does not mean that
the observed values accord with the predictions, it merely means that the null hypothesis has not been falsified. This may require more elaborate analysis or, as Ward
suggests, simply larger sample sizes. Unfortunately, even failure to conform to the predictions gives no information on which of the preconditions have been violated,
unless the data have been collected in such a way as to test a single, specific hypothesis for the population studied. And this is rare. For example, where Littorina spp.
diverge from HardyWeinberg equilibrium they tend towards excess homozygosity (Ward 1990), as do many other marine invertebrates (e.g. Crisp 1978, Singh &
Green 1984). This could be accounted for by a number of hypotheses including differential selection between larvae and adults (Crisp 1978, Zouros & Foltz 1984) and
the Wahlund effect. This is an apparent excess of homozygotes if it is incorrectly assumed that a series of local populations is a single, large, randomly mating
population. There is also evidence from several species for preferential survival of homozygous individuals on exposure to various forms of pollution (Lavie & Nevo
1986, 1987, Fevolden & Garner 1987), although there is no correlation between heterozygosity and either growth or oxygen consumption under starved conditions in
L. littorea (Foltz et al. 1993).
One selectionist theory predicts that, within a particular taxon, the degree of population heterozygosity will be affected by niche width, with a positive correlation
between niche breadth and genetic diversity (Van Valen 1965). A major difficulty with this theory is the definition of niche breadth. This is frequently assumed to equate
with the geographical range of a species or the number of habitats it occupies. Noy et al. (1987) found a correlation between niche width and heterozygosity in a
comparison of Nodilittorina punctata and Melarhaphe neritoides (as Littorina punctata and L. neritoides) although niche was defined rather loosely, and Janson
(1985b) found a correlation between heterozygosity and distribution of three Littoraria species from the Caribbean. Beardmore & Morris (1978) provide indirect, but
more convincing, support by showing that Littorina littorea had reduced heterozygosity where it cooccurred with L. saxatilis (they used the name L. rudis). The
implication is that cooccurrence leads to a reduction in niche breadth.
Apart from selective processes, heterozygosity within populations may also be affected by unpredictable interruptions of gene flow. Because of the nature of the
phenomenon and the difficulty of experimental work, papers concerning the effects of random processes such as genetic bottlenecks on heterozygosity are often highly
speculative. Nevertheless, there are convincing examples of the profound effect of such events on the heterozygosity of at least one nondispersive species. Populations
of L. saxatilis are divided into almost discrete breeding units (Janson & Ward 1984) and, with limited migration (Janson 1983) and brooding of crawling young, there
is very little gene flow between populations. This results in measurable genotypic variation on a scale of metres (B.Johannesson & Johannesson 1990) making this an
ideal candidate for stochastic effects. New populations established by a small number of individuals (theoretically a single pregnant female would suffice) may
experience a genetic bottleneck through a loss of genetic variability, i.e. a founder effect. This may be ameliorated by attributes of the population. Nei et al. (1975)
modelled the effects of a population passing through a genetic bottleneck, such as a founder event. They found that the number of alleles at each locus is profoundly
affected by the “size” of the bottleneck, but the reduction in average heterozygosity depends very strongly on the rate of subsequent population expansion. Janson
(1987b) and Johannesson & Warmoes (1990) suggest that rapid
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population growth accounts for the relatively minor loss of heterozygosity in L. saxatilis populations on isolated skerries in Sweden and breakwaters in Belgium,
despite presumed founder events. Johannesson & Warmoes (1990) also cite lack of correlation between the age of manmade breakwaters and heterozygosity as
implying lack of gene flow. However, very high gene flow would produce the same result by raising populations of different ages to the same level of heterozygosity.
This is more consistent with the rapid colonization observed, as it implies frequent recruitment from other populations.
Founder effects come out more clearly when more isolated populations are considered. Venetian populations of L. saxatilis may have been introduced by man
(Smith 1982) and show a markedly lower heterozygosity than British and Swedish populations (Janson 1985a). But the most vivid example of founder effects is shown
by two very localized and geographically separated populations of L. saxatilis in South Africa. Kilburn (1972) identifies fossil littorinids from South Africa as L.
saxatilis. But identification of fossil littorinids is difficult and Hughes (1979) and Knight et al. (1987) consider L. saxatilis to be an exotic species introduced to two
natural harbours by early European shipping. They show that within population heterozygosity is dramatically lower even than for L. saxatilis from Venice. They also
record a greater genetic distance between the two South African populations than among North Atlantic populations. Perhaps most striking is that both Venetian and
South African populations are genetically more distant from North Atlantic L. saxatilis than North Atlantic L. saxatilis are from British L. arcana.
Balanced against processes that reduce heterozygosity within populations are factors that promote gene flow among populations. Adult littorinids are normally
assumed to disperse over very limited distances (e.g. Smith & Newell 1955, Hamilton 1978, McQuaid 1981, Petraitis 1982, Janson 1983), although this is not always
so (see below). Assuming low adult dispersal, gene flow depends on larval dispersal. In this case, the production of crawling larvae by ovoviviparous species
presumably results in much less gene flow than among populations of species with planktonic larvae. It, should be possible, therefore, to make predictions about genetic
diversity on the basis of developmental mode. All species may be subject to localized selective forces that differ between populations, but for species with non
dispersive larvae there are the additional problems of inbreeding, genetic bottlenecks, founder effects, etc. Both neutral and selectionist theories predict that genetic
variability within populations should be reduced for nondispersive species, while differentiation among populations should be greater.
Burton (1983) reviewed data for natural populations of marine invertebrates and found that there is little evidence to support these predictions. Species with high
capabilities of dispersal often have substantial differentiation, either because of local selection or because of restricted gene flow between populations. Among the
littorinids, the evidence is contradictory and exists at several different levels. First I shall consider the evidence in support of these predictions.
Evidence in support of the theory

Some species with dispersive larvae are among the most variable morphologically, but this is normally withinpopulation polymorphism, in other words, each population
tends to be variable. Species with nondispersive larvae often have a high degree of variability amongpopulations, that is polytypic or geographic variability (Snyder &
Gooch 1973, Reimchen 1981, Naylor & Begon 1982). Likewise, some earlier studies on allozymes found that predictions about heterozygosity based on development
did apply (e.g. WiumAndersen
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1970, although she found both among and withinpopulation heterozygosity to be higher for a nondispersing species, Berger 1973, Snyder & Gooch 1973). Ward
(1990) compared data on heterozygosity for populations of seven species that had been examined for ten or more loci. He concluded that, on the whole, mean genetic
differentiation among populations for species with pelagic larvae was indeed lower than for species with crawling larvae. Likewise, Johannesson (1992) found little
amongpopulation variation over thousands of kilometres in populations of L. littorea and Melarhaphe neritoides. Liu et al. (1991) found the same for the thaiidid
snail Stramonita haemostoma; all of these species have dispersive, planktonic larvae.
There is also indirect support for these predictions in comparisons of brooders with species laying benthic egg masses. Species that brood or that lay eggs on algae
(e.g. Littorina mariae and L. obtusata) should have greater gene flow through dispersal by floating or rafting (Highsmith 1985, Martel & Chia 1991a,b) than species
laying egg masses directly onto the rocks. This applies in the case of the viviparous brooder L. saxatilis that does have greater withinpopulation heterozygosity than
sympatric L. arcana, which lays benthic egg masses (Ward & Warwick 1980, Ward & Janson 1985). However, it does not apply to the species examined by Knight
& Ward (1991). They found that the primary difference was between the oviparous L. nigrolineata and two species with different types of development, L. saxatilis
and L. arcana. L. nigrolineata is rarer and these results may reflect the size of the breeding populations.
There is evidence that poorly dispersed species are more locally adapted. For example, the phenotypic response shown by New England L. obtusata (but not L.
littorea) to the introduction of crabs, described above. There is also experimental evidence. When collected from different sites and translocated onto the same shores,
individuals from different L. littorea populations grew at similar rates, whereas L. saxatilis growth depended on the original site of collection (Behrens Yamada 1987).
Behrens Yamada suggests that L. littorea achieves a broad distribution through individual plasticity, while L. saxatilis does so by means of locally adapted ecotypes
(e.g. Grassle & Grassle 1978). She has also shown the same effect in a comparison of two Pacific species (Behrens Yamada 1989).
Finally, there is support for the theory from studies of speciation. Vermeij (1972) notes that highshore species tend to be both less dispersive and more endemic
than subtidal species. However, he suggests that these are independent responses to the same factors, with no cause and effect relationship. If there is a link between
larval development and gene flow, then developmental mode should influence rates of speciation. Reduced gene flow may increase the potential for rapid evolutionary
responses to changes in selective forces. We have seen this in the shell thickening of North American L. obtusata (Seeley 1986) since the introduction of shellcrushing
crabs, although this may be confounded with phenotypic responses. The effect of gene flow on evolution depends on the stability of population structure over
evolutionarily significant time (Slatkin 1987) and nondispersive species have all the genetic prerequisites for rapid speciation (Crisp 1978, Scheltema 1978, Jablonski
& Lutz 1983, Jablonski 1986). There is some evidence to support this.
The absence of dispersive larvae may enhance speciation because of stochastic effects such as greater amongpopulation heterozygosity caused by founder effects
and genetic drift. For example, Reid (1990a,b) believes that at least two species of Littorina may have colonized the North Atlantic from the North Pacific. One of
these led to the monospecific L. littorea clade, which has dispersive larvae, while the other has rapidly radiated into a clade of six species with nonplanktonic
development. This hypothesis is supported by an examination of the allozyme similarities of Pacific and Atlantic species by Zaslavskaya et al. (1992).
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Evidence against the theory

Other studies, either directly or indirectly, fail to support the predictions. Although direct developers can show geographic variability, the degree of morphological
variability within a population can differ between sites. According to Ockelmann & Nielsen (1981), the highly dispersive Lacuna vincta is polymorphic throughout
Europe, but morphological variability in the direct developers L. parva and L. pallidula is populationdependent. Polymorphic populations occur in Britain and France
while Danish populations of both are monomorphic. On the molecular level, Wilkins & O’Regan (1980) found that the relationship holds between amongpopulation
heterozygosity and reproductive mode, but is weaker than previously observed. Gaines et al. (1974), examining a single locus, found significant genetic differences
among populations of Littoraria angulifera (as Littorina angulifera) on islands that were separated by as little as hundreds of metres, although this species has
planktotrophic larvae. Kartavtsev & Ephremov (1981) found significant differences in heterozygosity of L. brevicula and L. mandshurica (which is nearly
monomorphic), even though both have planktotrophic larvae.
We must also consider Ward’s (1990) data more carefully. The data he summarized show that there is no significant difference in average withinpopulation
heterozygosity between dispersive and nondispersive developers (hs in Ward’s table 2; ttest, P>0.05). He does note higher amongpopulation genetic differentiation
for species with nondispersive larvae, but the highest value shown is in a comparison of “worldwide” L. saxatilis populations. These include data for Italian, North
American and South African populations. It is unreasonable to include this information as we have seen that there is evidence for the effects of genetic drift on several of
these populations. All other comparisons are for populations that are, at most, hundreds of kilometres apart. If this single datum is omitted, a ttest shows no significant
difference in minimum genetic distance among populations of species of the two developmental types (dm on Ward’s table 2; P>0.05). More recently, Tatarenkov
(1992) found no significant difference in the mean withinpopulation heterozygosity for seven species with pelagic development and ten without a pelagic stage.
Janson’s (1987a) results also deviate from some of the predictions. Like WiumAnderson (1970), she found higher withinpopulation heterozygosity in L. saxatilis
(which has crawling larvae) than L. littorea (pelagic larvae). This could be interpreted in terms of Ohta’s (1974) theory, assuming a smaller effective population size for
L. saxatilis. Janson also found no correlation between reproductive mode and overall allozyme heterozygosity in a comparison of published data for ten species. She
concludes that the dispersive ability of larvae influences gene flow, but that other factors, such as historical events, may be more important in determining genetic
variability within populations.
Genetic variability is influenced by factors other than gene flow and a major difficulty with interpreting some of these results is that genetic variability is measured in
adults, that is successful recruits. We know nothing about selection pressures in the plankton, or immediately after settlement. It is assumed, on the basis of recruitment
data, that mortality of planktonic larvae is very high (e.g. Underwood & McFadyen 1983, McQuaid 1992) and densities of newly settled larvae may be diminished by
>90% in the first 24h after settlement (Fretter & Manly 1977). I have not found estimates in the literature of mortality rates for newly emerged direct developers. New
born L. rudis (=L. saxatilis) are unexpectedly hardy (Berry & Hunt 1980), but their mortality is also high (e.g. Hughes & Roberts 1981). Survival of either planktonic
or newly settled or emerged juveniles may be genotypedependent. Ward (1990) found no evidence for genotypedependent survival of newborn young in breeding
experiments
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with L. saxatilis, but survival is genotypedependent in larval Mytilus edulis (Mallet et al. 1985). Withinpopulation heterozygosity would be influenced by, for
example, selection favouring different homozygotes in the larval and adult stages (Crisp 1978).
A final anomaly is offered by amongpopulation variances in heterozygosity of Bembicium vittatum, which has little or no pelagic development (Reid 1988). This
species occurs in lagoons as well as on the open coast. One would expect better mixing of the gene pool in lagoons, yet, over similar distances, variance between
populations is greater in lagoons than on exposed shores (Johnson & Black 1991).
We are left with the paradox that there is an imperfect correlation between mode of development and heterozygosity and yet species with nondispersive larvae
generally show more geographic variability, and stronger local adaptation. Part of the problem may be the link between molecular and morphological characters.
Another problem may lie in assuming that valid generalizations exist. Like shell polymorphism, heterozygosity is influenced by many factors and the balance may depend
on either the species involved, or even the population. Littorina littorea is thought to be exotic to North America (see McQuaid 1996) and Berger (1977) suggests
that initial colonizers went through a bottleneck that produced very low variability in the population he studied. He found a large genetic distance between L. littorea
from the USA and France, but Morris (1979, cited in Ward 1990) did not find this in a comparison of L. littorea from the USA and Wales. Certainly there may be
something odd about L. littorea. While L. littorea and Melarhaphe neritoides both conform to the predictions on amongpopulation variation for dispersive species,
the former showed much lower withinpopulation variation than predicted by either selectionist or neutral theory (Janson 1987a, Johannesson 1992).
In summary, the evidence for the effects of development mode on both within and amongpopulation heterozygosity is conflicting. Part of the difficulty is that these
studies concern correlations between the two factors. While development mode is fixed, heterozygosity within a population is affected by different factors in different
localities, and probably at different times. Even assuming that initial recruitment of juveniles is not influenced by their genotype, with a wide range of species occupying
many sites we cannot exclude the probability of different combinations of selective forces operating on each population. Amongpopulation variability in part reflects the
degree to which these forces produce different results in each population. The fact that there is no clearcut correlation between development and heterozygosity
suggests that development may influence heterozygosity, but does not have an overriding effect. Other factors, or combinations of factors, may be more important in
some populations or at different times.
Dispersal
Dispersal has both ecological implications (recruitment) and evolutionary implications (gene flow) and dispersal ranges are expected to correlate with the mode of larval
development. Pelagic larvae are capable of dispersal over long distances, but the distances involved depend on local current systems, the longevity of the pelagic phase
and also larval behaviour. The planktonic diatom Anaulis australis, is permanently retained within the surf zone off sandy shores by a combination of longshore gyres
and diurnal vertical migration patterns that take advantage of counter currents at different depths in the water column (Talbot et al. 1990). At the other extreme,
Scheltema (1971) estimated that 60 days in the water column was sufficient for currents to transport larvae across the Atlantic. On the basis of this, Reid (1986)
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suggests that lack of morphological differences between Littoraria angulifera on either side of the Atlantic could be due to larval exchange. The dearth of information
on the actual range of dispersal of littorinid larvae is in accord with our general lack of understanding of events between spawning and settlement of benthic species.
Recent work on a number of marine taxa suggests that the link between dispersal and mode of development is not always what we might assume. Direct developers
seem to be capable of greater dispersal than expected through rafting or drifting of egg masses or postlarvae and postmetamorphic drifting may be quite common
among both gastropods and bivalves (Highsmith 1985, 6 Foighil 1989, Martel & Chia 1991a). In the case of the bivalve Gaimardia, dispersal on kelp rafts occurs
over hundreds of kilometres (Helmuth et al. 1994). Lacuna is known, particularly under unfavourable conditions, to promote drifting actively by producing mucous
threads and lifting the foot and is capable of at least smallscale dispersal in the water column as adults (Johnson & Mann 1986, Martel & Chia 1991a,b, Martel &
Diefenbach 1993).
On the other hand, correlations between adult and recruit density of some species indirectly suggest that the dispersal of planktonic larvae may be limited under some
conditions (e.g. Hunte & Younglao 1988, Meekan et al. 1993, Peterson & Summerson 1992).
Feeding larval forms are thought to be both ancient and conservative (Strathmann 1985). But, while possession of a feeding larva may have profound effects on
dispersal (and hence range, local adaptation, rates of speciation etc), it has been argued that it is not dispersal itself which is selected for (Palmer & Strathmann 1981,
Strathmann 1985). Rather, the primary advantages of a planktonic larva are associated with feeding and the advantage of planktotrophy over lecithotrophy lies in
aspects of life history such as the ability to produce more, smaller eggs.
Scheltema (1989) found that, among a number of prosobranchs, species with planktonic larvae had a greater mean geographical range than those with non
planktonic development. But among the littorinids, some observations of actual dispersal and range of species with different types of larvae are distinctly counter
intuitive. Within the genus Littoraria, L. aberrans is unique in being ovoviviparous with crawling larvae and, as expected, it has a very restricted range (Reid 1989).
Yet L. albicans, which appears to have a particularly protracted planktotrophic life, also has a limited range (Reid 1986). This could be due to very specific habitat
requirements or narrow tolerance of physical conditions in the case of L. albicans, but other anomalies cannot be easily explained. Johannesson & Warmoes (1990)
found that Littorina saxatilis, which lacks a planktonic stage, colonized manmade breakwaters on the sandy Belgian coast almost as efficiently as L. littorea.
Oceanic islands often support species exhibiting direct development. A tropical example is given by Rosewater (1982). Other examples among the littorinids are
given by Simpson (1977), Picken (1979) and Simpson & Harrington (1985), but these are from the Southern Ocean and this adaptation could be linked to sub
Antarctic conditions. Most benthic polar invertebrates are direct developers, possibly because of the poverty or unreliability of phytoplankton food supplies (Thorson
1950, Chia 1970, Mileikovsky 1971). Limited dispersal of oceanic species is in accord with the predictions of the theory of island biogeography (MacArthur & Wilson
1967) and is presumed to be safer than producing planktonic larvae that will be carried away by oceanic currents with little chance of recruitment back into the parent
population. But there is also the unexpected finding that, even in a relatively restricted sea such as the North Atlantic, L. saxatilis has a more widespread distribution
than L. littorea (Johannesson 1988). Not only is this contrary to expectations based on mode of reproduction, but it also runs contrary to a prediction made by
Vermeij (1972) based on the
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physical uniqueness of habitats. He argues that low intertidal environments are more uniform than the upper shore and, therefore, that highintertidal species would have
narrower geographic ranges than lowshore species. L. saxatilis generally occurs higher upshore than L. littorea and yet is more widespread. It is possible that L.
littorea is indeed more widely dispersed than L. saxatilis, but less tolerant of the range of abiotic environments it encounters upon, or during, dispersal. Obviously our
understanding of the control of species’ ranges and dispersal is not what it might be.
Life history theory

Life histories have both quantitative aspects (such as size at maturity, number of eggs produced, etc.) and qualitative aspects such as the mode of larval development.
Quantitative aspects are relatively plastic and are under local selective pressures, resulting in differences among populations. An emerging view is that qualitative aspects
are largely fixed through phylogeny and changes require evolutionary or speciation events.
The reproductive effort of gastropods depends on factors such as body size (e.g. Grahame 1973, Hughes & Roberts 1980, Underwood & McFadyen 1983), and
food availability (e.g. Hughes 1971, Spight & Emlen 1976). However, reproductive patterns within the same species can also vary with habitat. L. saxatilis shows the
same trends between exposed and sheltered rocky shores as between sandy shores and rocky shores. In both comparisons the first given habitat is associated with
smaller body size, earlier maturity, higher reproductive effort and more numerous, smaller embryos (FallerFritsch 1977, Atkinson & Newbury 1984, Gallardo &
Götting 1985). This is seen to be related to habitat stability and the probability of adult or juvenile mortality. Selective predation of specific size classes has also been
speculatively suggested as an explanation for early maturity of “dwarf” populations of L. mariae (Reimchen 1982).
Theories of life history attempt to provide generalized predictions on the reproductive strategies exhibited by organisms depending on the circumstances under which
they live. This is only possible if the intrinsic (physiological) tradeoffs or constraints are understood and the demographic effects of extrinsic (ecological) factors
recognized (Calow 1983). With this proviso in mind, field evidence frequently suggests correlations between environmental patterns and trends in the mode of
reproduction. This is seen to be the evolutionary result of balancing costs and benefits in the process of adaptation where the major constraints on strategies are the
spatial and temporal heterogeneity of the habitat (Southwood 1977, 1988). There are two approaches, rK selection and “bethedging”, that are summarized by
Stearns (1976) and compared with available data by Stearns (1977). The predictions of these theories depend largely on habitat homogeneity (in time and space),
which is reflected in relative mortality rates of adults and juveniles. rK selection predicts a number of linked reproductive traits associated with stable and unstable
environments, defining stability in terms of adult survival. Bethedging has similar predictions for situations in which adult mortality is variable, but predicts different
combinations of traits when juvenile survival is variable.
Littorinids are convenient organisms for testing such theories because of the wide range of reproductive strategies encountered among them and the wide range of
habitats they occupy. A number of authors have surveyed the occurrence of particular modes of larval development among the Littorinidae and correlated them with
habitat, particularly zonation (e.g. Woodward 1909, Anderson 1961). These correlations have been imperfect and unsatisfying (Mileikovsky 1975, Reid 1989). More
detailed studies have been made of predictions concerning differences among species and also within a single species living in different
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habitats. On the whole, these studies have brought out even more strongly the lack of fit between predicted and actual strategies of littorinids.
Grahame (1977) compared two semelparous Lacuna spp. (i.e. species that breed once in their lifetimes). These were L. vincta (planktotrophic larvae) and L.
pallidula (direct, lecithotrophic development). Contrary to the theoretical predictions, reproductive effort was twice as large in L. vincta. Grahame explains this by
considering L. vincta to be more rselected. Although L. vincta does have survival curves that indicate heavier mortality of juveniles, both species live for only about
one year and are towards the r end of the rK continuum (Smith 1973). Subsequently, Grahame (1982) calculated reproductive effort in energetic rather than biomass
terms and concluded that the difference between the two species was minimal. This emphasizes the importance of applying the correct measure of reproductive effort.
Hughes & Roberts (1980, 1981) compared two directdeveloping Littorina spp. with two species showing planktonic development. Although effort (estimated as
energy equivalent from biomass) per individual juvenile was greater in the direct developers there was no correlation between the mode of reproduction and either total
reproductive effort or adult mortality rates. They did find differences in egg size and reproductive effort among populations of L. rudis in different habitats, but these did
not correlate with demographic differences. This was taken further by Hart & Begon (1982) who compared cliff and boulder populations of L. rudis. They too found
the theories inadequate, as their populations exhibited unpredicted combinations of traits (i.e. a mixture of r and K traits). This was explained by local variation in
factors influencing shell thickness and animal size (crab predation and wave action), causing Hart & Begon to question the value of generalized theories.
Robertson & Mann (1982) studied the life history of an animal they called L. neglecta living in Zostera meadows. They found very high mortality of recruits in late
summer and autumn. Size specific fecundity was much higher than for L. saxatilis while growth, size at maturity, egg weight and size at hatching were all much lower
(cf. Berry 1961, Hughes & Roberts 1980, 1981). Reproduction was thought to account for most of the energy used by these populations, suggesting selection for life
history traits linked to high juvenile mortality. Recently B.Johannesson & Johannesson (1990) found that the populations of L. neglecta studied by Robertson & Mann
are actually ecomorphs of L. saxatilis. If this is correct then it may represent a case of life history adaptation within a single species when more extreme selection is
applied than in the studies of Hughes & Roberts (1980, 1981) and Hart & Begon (1982). The fact that L. neglecta is thought to be semelparous, unlike L. saxatilis
(HannafordEllis 1983, 1984, Fish & Sharp 1985), may just reflect the probability of individual survival. The populations of L. saxatilis Robertson & Mann studied in
Zostera meadows suffer intense densitydependent mortality among recruits and 2yearold animals. Like the very slow growth rates measured, this is presumably due
to food shortage (Robertson & Mann 1982). There is a low probability of any individual reaching 3 years of age so that the population will be effectively semelparous,
even if longer lived individuals were capable of breeding repeatedly.
The difficulty of applying life history theory to comparisons of littorinid species may be resolved by considering evolutionary constraints. Stearns (1976) points out
that selection does not necessarily produce the optimum solution to a problem, but the best available solution. Potential phylogenetic constraints are recognized at higher
taxonomic levels. For example it is suggested that the nereid polychaetes and the cephalopod molluscs have lost the genetic potential for iteroparity (Olive 1985). The
choice between iteroparous and semelparous reproduction is likely to come under such constraint as each path involves major physiological commitments in the adult.
We might expect less marked evolutionary
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constraints in the choice between planktotrophic and lecithotrophic development. This distinction is ecologically critical to larvae, but less so to adults as it does not
require major differences in physiological adaptation. Brooding may require morphological changes such as differences in shell shape depending on brood size (e.g.
Grahame & Mill 1992), but the gonadotrophic hormone/spawning hormone system is versatile enough not to dictate the use of particular forms of iteroparous
reproduction (Olive 1985). For example, it has been suggested that Littorina arcana is a reproductive morph of L. saxatilis (Caugant & Bergerard 1980, Bergerard
& Caugant 1981, Hughes & Roberts 1981) although this view is not widely held.
Reid (1989, 1990b) argues convincingly that differences in mode of development of cooccurring littorinids do not reflect ecological differences, but phylogenetic
differences. The taxonomic clades that he recognizes are characterized by, among other things, their mode of development. Indeed this forms one of the characters on
which the analysis is based, but he avoids circularity by obtaining the same phylogenetic tree using an analysis that omits this character.
Phylogeny can resolve seemingly intractable ecological problems. But it does more than this by illustrating that even logical ecological explanations can be spurious,
unless framed against a phylogenetic background. As an example, the need to explain differences in the mode of reproduction of Melarhaphe cincta and M. scabra on
an ecological basis falls away with the reclassification of these species. M. cincta is now classified as Nodilittorina cincta (this genus has pelagic egg capsules) and
M. scabra as Littoraria scabra, falling into a subgenus with ovoviviparous development (Pilkington 1971, Reid 1986, 1989). Ecological explanations remain valid
where a species shows an adaptive, derived condition that differs from the primitive condition for the clade to which it belongs.
Reid (1991) gives examples from the subfamily Littorininae, which are mostly littoral fringe species. While benthic egg masses are primitive in the family Littorinidae
as a whole, this subfamily produces pelagic egg cases. Benthic egg masses can be vulnerable to desiccation and Reid suggests that pelagic egg cases are advantageous
to species living high on the shore. Thus, within this subfamily planktonic development is the primitive state. Direct development in members of this subfamily is a
derived condition that may then be explained as an adaptation to particular ecological conditions.
Conclusions
There are areas of confusion within both major topics covered in this review; morphological variability and larval development. Considering morphological variability,
there seems to be a contradiction between ecological and evolutionary perceptions about the importance of predation. Predation is frequently cited as a driving
mechanism for supposedly adaptive morphological differences between coexisting species, or between conspecific populations living under different predator regimes
(e.g. Boulding & van Alstyne 1993, literature cited above). This includes intraspecific morphological differences between sites with different degrees of wave action as
wave action can influence predator abundance. The problem is that ecological papers that deal directly with rates of predation in the field almost all find that predation is
not important at the population level (see references in McQuaid 1996).
While we must accept that field observations may underestimate predation, many ecological studies specifically comment on either low rates of predation (e.g.
Johnson & Mann
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1986, Huxham et al. 1993), or the fact that the apparently high numbers of littorinids eaten by predators represent a small proportion of the total population (e.g.
McQuaid 1985, Alerstam et al. 1992). Although local densities of littorinids may reflect the presence of predators, there is no evidence of littorinid populations being
limited by predation.
There are several ways of resolving the views of ecologists and evolutionary biologists on the importance of predation. First, predation may be intense enough to
drive evolution, without having a marked effect at the population level.
Secondly, there is the rather damning comment that evolutionary studies that invoke predation to explain morphological variability are largely based on qualitative
correlations and do not actually measure predation. Even those evolutionary studies that demonstrate differential survival of contrasted morphs or species rely on
laboratory experiments, use prey that are restrained in the field and so overestimate predation rates, or ascribe differential survival in the field to unknown, hypothetical
predators (e.g. Seeley 1986, Reid 1987, Reimchen 1989).
Thirdly, it is possible that levels of predation are so low and gene flow so high (even for direct developers) that polymorphism reflects phenotypic phenomena rather
than genotypic. Two examples show how alternative explanations can be sought for polymorphic phenomena. The sandy beach bivalve Donax serra is highly
polymorphic in South Africa and this could be ascribed to reflexive evolution; high intraspecific variability preventing visual predators from developing strong search
images. Yet this is nonsensical as the animal is permanently buried and so invisible to predators. Likewise, in some littorinids shell shape is more “predator resistant” in
areas where there are more predators. An explanation that does not require genotypic responses is that shell shape depends on growth rates that differ among sites and
which may also be phenotypically influenced by changes in the behaviour of snails in the presence of predators (Appleton & Palmer 1988, Palmer 1990).
Finally, ecologists and evolutionists tend to study different circumstances. Evolutionary studies are often concerned with polymorphic (or polytypic) situations. In
some cases at least polymorphism and the existence of cryptic species make ecological work impossible. Evolutionary studies may address situations where predation
is high enough to drive polymorphisms and ecological studies address situations where predation is not strong enough to drive polymorphisms.
There is a similar problem with convergence of form in several species in the same environment and of widely separated populations of the same species assuming
similar morphologies in similar habitats. Is this genotypic, or is it a phenotypic consequence of some environmental influence on growth rates?
We can begin to separate these hypotheses by doing ecological studies of natural predation rates in polymorphic populations to determine how intense natural
predation rates actually are and whether they really are morph specific (e.g. studies of predator gut content studies). The most effective way to resolve the question of
whether habitatdependent polymorphism (i.e. polytypy) has a genotypic or phototypic basis is to do breeding experiments.
On the topic of larval development, littorinids do not conform well to the predictions of life history theory. Correlations between mode of development and vertical
zonation are poor and studies of differences among species and also within a single species living in different habitats point to a gap between theoretical predictions and
what littorinids actually do. A solution to this dilemma may lie in the evolutionary history of taxa as this constrains the type of development they can exhibit. Predictions
of life history theory emphasize ecological differences among populations, but need to take cognisance of phylogenetic constraint.
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Likewise the link between type of larval development and heterozygosity is poor. In part this reflects the fact that heterozygosity (unlike mode of development) is not
fixed and responds to a wide variety of factors. The search for a general correlation between larval development and heterozygosity may be misleading. Factors that
influence heterozygosity include gene flow, which may be influenced by larval dispersal, but other effects may be more important in some populations. The problem is
exacerbated by the lack of correlation, in at least some cases, between mode of development and the extent of larval dispersal. This is important as larval dispersal is
central to evolutionary studies since it influences gene flow, and to ecological studies as it affects recruitment (see McQuaid, 1996).
Acknowledgements
It is a pleasure to acknowledge the various sources of funding for this work. The work was begun during the tenure of an Ernest Oppenheimer Memorial Trust Fund
Travelling Fellowship and further funding was provided by the Joint Research Committee of Rhodes University and the Foundation for Research Development. I am
very grateful to A.D.Ansell for his patience as an editor and to the people who took the time and effort to comment on earlier drafts of the paper: A.R.Beaumont,
R.J.Chambers, P.E.Hulley, D.G.Reid and especially K. and B.Johannesson. My thanks also go to Lesley McQuaid for coping with the references.
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BIOLOGY OF THE GASTROPOD FAMILY LITTORINIDAE.

II. ROLE IN THE ECOLOGY OF INTERTIDAL AND SHALLOW MARINE
ECOSYSTEMS
CHRISTOPHER D.McQUAID
Department of Zoology and Entomology, Rhodes University,
Grahamstown 6140, South Africa

Abstract Littorinid gastropods are cosmopolitan and ubiquitous in intertidal and shallow marine ecosystems. In many cases they have a profound
influence on the structure of these systems. There is an extensive literature on the ecology of littorinids, dealing in greater or lesser detail with the
autecology of a large number of species. This paper summarizes the literature on community interactions, concentrating primarily on the best studied
examples of the main feeding types. The effects of predators are described for each case. Although predation is often cited as central to selective
processes in the literature on the evolutionary biology of littorinids, in most of the ecological literature predation is not seen as controlling population
densities.
Individual species of littorinids can feed on many different types of food, but most fall reasonably well into one of four categories. These are:
detritivores (that eat dead angiosperm material); epiphytic grazers (that eat the periphyton of algae and angiosperms); macroalgal grazers; and epilithic
grazers (that brush microalgae and sporelings from the rocks). Community interactions can be complex and are extremely variable from place to place
and time to time, but it is possible to make generalizations. The effects of littorinids on community structure depend primarily on feeding type. Species
with strong direct effects on the distribution and abundance of other organisms fall into the last two categories, species in the first two categories have
weak direct effects.
Neither detritivores nor epiphytic grazers have been shown to influence community structure strongly. Detritivores have no recorded effect on
species densities or distributions, but can be very important in nutrient cycling in saltmarshes. They eat both angiosperm detritus and the fungi
colonizing dead plants, making refractory angiosperm material available as faeces to other organisms. Some, for example Littoraria irrorata, are
important prey for predators such as conchs and crabs and provide hermit crabs with shells, that can control their population structure. Epiphytic
feeders have no direct effect on macroalgae. Several authors have commented on the negative effects of epiphytic plants on macroalgae (increased
shading, drag and frond breakage) and it has been suggested that epiphytic grazers may indirectly influence community structure by removing epiphytes
and enhancing the fitness of their host plants.
The situation can be quite different for the remaining two feeding types. Littorinid macroalgal grazers mostly occur on the low shore or subtidal and
can control algal biomass directly. While macroalgal grazers of other taxa have been found to influence algal zonation, it it is not clear that littorinids do
so. Feeding behaviour and preferences reflect algal defences, both chemical and physical. Both types of defence deter generalist grazers, but chemical
defences can be overcome by specialist grazers that are immune to the chemicals produced by a particular alga. The amount of damage that
macroalgal grazers do depends not so much on grazer density as on fine details of the interactions between grazer and alga pairs. A grazer may
devastate one species of alga, but do little damage to another, even at similar grazer densities.
Lastly, epilithic grazers have the most dramatic influence on community structure. Whereas macroalgal grazers affect algae by reducing biomass or
survival of mature plants, this group can control the recruitment of algae. Epilithic grazers occur throughout the shore and their effects vary with height
on the shore. This is because the consequences of grazing depend not only on grazing pressure, but also on algal growth rates and algal competitive
hierarchies. Because the effects of epilithic grazers depend on the balance between grazing and algal growth or recruitment, they are chiefly density
dependent. Grazing effects operate mainly through the removal of macroalgal sporelings so that mature plants represent sporelings that have “escaped”
grazing. These grazers may exclude mature macroalgae where particularly dense (as on sheltered shores), or
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have no appreciable effect where algal settlement or growth are particularly high. Epilithic grazers may also eat ephemeral algae. Whether their grazing
increases or decreases algal species richness depends on the position of their preferred food in the algal competitive hierarchy.
The primary reason for the complexity of these interactions is the fact that littorinid grazing rates, algal growth rates and algal competitive hierarchies
are all influenced by the same factors: height on the shore, wave exposure, habitat and season, i.e. grazers and algae react to the same factors and
interact among each other.
Epilithic grazers can also affect sedentary animals very strongly. The complexity and importance of interactions are clear when we consider epilithic
grazers in combination with either algae or barnacles. Algae or barnacles can be dense enough to inhibit grazers (either directly, or indirectly by
harbouring predators), or grazers can control recruitment of algae and barnacles. The direction of the interaction depends on the relative densities of
the species involved. As densities largely reflect recruitment success, a good understanding of recruitment processes is essential if we are to be able to
predict the results of species interactions.
Introduction
In a companion paper (McQuaid, 1996) I consider evolutionary aspects of the biology of littorinids, including shell polymorphisms (that may be affected by predation)
and the theoretical link between mode of larval development, extent of larval dispersal and gene flow. This paper concentrates on littorinid ecology. Specifically, I
discuss the effects of littorinids on communities while ignoring the many other aspects of their ecology that have been extensively studied. For example there are
substantial bodies of literature on population dynamics and shore height variations in population size structure (Raffaelli 1982), the effects of parasites (Lauckner 1984)
and the effects of pollutants, including the use of Littorina littorea and other common species as sentinel organisms (Bryan et al. 1983, Nevo et al. 1987). Here I
focus on species interactions between littorinids and algae, other grazers, predators, etc.
Phylogeny and ecology

Reid’s (1989) phylogeny of the Littorinidae suggests a close correspondence between ecological habit and phylogeny. For example, vegetationdwelling species fall
largely into the genus Littoraria, and Nodilittorina spp. are high shore grazers of microalgae and cyanobacteria in tropical and subtropical areas. This suggests that
phylogenetic lines at the level of subfamily, or thereabouts, represent lines of ecological experimentation, followed by radiation within particular habitats or modes of life.
This is an intuitively attractive view. Allowing for ecological plasticity (e.g. Littorina littorea and L. saxatilis both occur on rocky shores and also in saltmarshes),
taxonomic groupings tend to coincide with broad ecological habitats (e.g. dwelling on hard or soft substrata), and so with the ecological systems where they occur (e.g.
on rocky shores or on mangroves). This may reflect the need for general adaptations for particular habitats. Southwood (1977, 1988) explores the relationship
between habitat and reproductive strategy, and Reid (1989) suggests that reproductive mode or spawn type is one of the factors allowing colonization of the littoral
fringe by the Littorininae. But among the littorinids we also see convergence of form within the same habitat. For example L. littorea occurring in New England
saltmarshes tend to resemble the marsh snail Littoraria irrorata (Vermeij 1978) while there is, presumably adaptive, convergence of form in a number of
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European species of Littorina where they occur in the barnacle zone (Reid 1993). Speciation may result in the genetic fixation of an ecotype. Association with a
particular habitat also puts constraints on other aspects of an animal’s ecology, such as the way it feeds. Highshore species living beyond the physiological limits of
adult or sporeling macroalgae have little choice but to eat microalgae, while those living on angiosperms must either solve the problem of digesting refractory plant
material or feed on periphyton or detritus. In temperate areas, midand lowshore species as well as subtidal species often occur in areas dominated by dense stands of
macroalgae and tend to eat either macroalgae or algal epiphytes.
Grazing
Grazing by littorinids has recently been reviewed by Norton et al. (1990) who provide details on mechanisms of grazing, feeding behaviour, aspects of algal
attractiveness and algal defences against grazers. Briefly, the taenioglossan radula of littorinids is versatile enough to allow a wide range of both macro and microalgae
to be used as food. Differences in radula structure suggest a basis for resource partitioning, but it is not possible to predict diet from the radula (see also Hawkins et al.
1989). Some species, such as L. littorea and L. scutulata are opportunistic generalists while others such as L. obtusata have more specialized diets. However, even
generalist species can have distinct preferences among macroalgae. L. littorea differentiates between extracts from different algae, responding most positively to those
from ephemeral species (Imrie et al. 1989).
Littorine grazing can reduce the biomass of microalgae both in the laboratory and in the field (Castenholtz 1961, 1963, Nicotri 1977, Underwood 1984a, Williams
1992a) and selection for particular microalgae can drastically alter species composition of the microalgal assemblage (Hunter & RussellHunter 1983, Williams 1992a).
In contrast to macroalgae, selectivity is thought to be passive and determined by how tightly microalgal species adhere to the substratum and by their morphology.
Loosely attached diatoms and chain forming species are removed from the substratum more easily (Nicotri 1977), while smaller snails take smaller diatoms (Hylleberg
& Christensen 1978).
Community interactions
Underwood et al. (1983), Underwood (1985) and Underwood & Kennelly (1990) have clearly demonstrated the difficulties of defining community interactions as they
vary with both time and space, sometimes on very small scales. For example grazer/algal interactions are influenced not only by herbivore densities and feeding rates,
but also by algal recruitment and growth rates. So grazers may eliminate algae or be swamped by them depending on the height on the shore, season, between year
variations in weather and so on. Given this natural variability it is impossible, at this stage, to predict the outcome of interactions. In many cases we do not even know
what information we need to make accurate predictions. It is also difficult to compare the results of studies that were carried out with different degrees of resolution
(spatial or temporal) or rigour. Recognizing that species interactions and the effects of species on communities are not fixed, this paper reviews the available information
on the effects of littorinids on community structure.
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The first point is that these effects depend on what type of feeder a species is (e.g. Lubchenco & Gaines 1981). In the case of epilithic grazers particularly, their
influence on the community also depends partially on how high on the shore they live. Height on the shore influences not only attributes of the grazer, such as the
duration of foraging, but also attributes of the algae. These include settlement and growth rates, as well as competitive hierarchies among the algae. I have divided the
littorinids into four major feeding types.
(a) Epilithic grazers. These are species that use the radula primarily to scrape the surface of rocks taking in both microalgae, such as benthic diatoms, and the
microscopic sporelings of macroalgae. In several cases they can eat adult macroalgae as well, with a decreasing preference from microalgae ephemeral
macroalgae perennial macroalgae. They occur at all levels on rocky shores from high to low shore.
(b) Detritivores. These eat dead rather than live angiosperm material and may also eat the fungi colonizing plant detritus. These animals occur on angiosperms, such as
saltmarsh plants and mangroves. I exclude here species such as L. littorea, that feeds epilithically but will also eat drift algae.
(c) Epiphytic grazers. These feed on epiphytes on the surface of macroalgae or angiosperms. At least as adults they live more or less permanently on the surface of
macrophytes and are found on rocky shores, mangroves and in saltmarshes.
(d) Macroalgal grazers. These feed on mature plants, rather than sporelings and are mainly found on the mid to low shore and subtidally on rocky coasts.
Obviously a single species may feed on more than one type of plant material (for example L. littorea), but most species do not appear to be very flexible in their mode
of feeding and fall reasonably well into one of these categories.
Studies of the community effects of different littorinids are biased by the type of research done in the ecosystems where they occur. This reflects what is important, or
at least appears important, to ecologists working on these systems. On rocky shores there has been a great emphasis on the processes that control the abundance and
vertical distribution of species. These include the effects of disturbance, predation or grazing, and competition, which are often examined by using manipulative
experiments in the field. Littorinids from rocky shores tend to be studied from this perspective, with particular emphasis on their interactions with other species. In
contrast, research on saltmarshes tends to consider nutrient and energy flow with less manipulative experimentation, although this has increased recently.
There is scattered information on feeding and sometimes species interactions for a number of littorinids, but relatively few species have been studied in detail.
Fortunately, there is at least one well studied example of each of the four feeding types. I shall concentrate on the most extensively studied example of each, drawing in
information on other species where appropriate. Littorinids are eaten by a wide range of predators (reviewed by Pettitt 1975) which, in some cases, directly or
indirectly influence grazer density. But the effects of predators can vary enormously between species, or even populations of the same species. This is considered for
each of the examples given.
Epilithic grazers
Epilithic grazers can have a dramatic effect on the distribution and abundance of macroalgae through their effects on sporeling survival. Most feed on sporelings in
preference to mature plants and so sporelings that escape grazing can grow into a refuge in size. Because of this, the distribution of mature plants can reflect the
distribution of sporeling “escapes” from
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grazers. This emphasizes the importance of grazing pressure and of factors that promote the rate at which sporelings grow beyond the size at which epilithic grazers kill
them. Grazing pressure is a function of grazer density and behaviour. Density can be influenced by the presence of predators (either through loss of numbers from
predation or through predator avoidance) as well as the life histories and recruitment success of the grazers. It is also influenced by the availability of suitable
microhabitats, such as crevices, barnacles or algal canopy. Grazing pressure is influenced by several factors: (a) height on the shore, as most littorinids graze only when
submerged or damp (Little 1989), although littorinids with reduced foraging time may feed faster (Newell et al. 1971, Underwood & McFadyen 1983); (b) weather
(including season); (c) degree of wave exposure, as heavy wave action can inhibit feeding. Balanced against grazing pressure are sporeling recruitment and growth that
are also affected by several factors. Again, one of these is weather, including seasonality so that grazing pressure may be constant through the year but the effects of
grazing may be seasonal. A second factor is canopy of mature plants, which can influence several things. Algal canopy can promote sporeling survival in the absence of
grazers or it can shade out sporelings. Alternatively, algal canopy can promote aggregation of grazers so that local densities of grazers are greater. A third factor
influencing sporeling growth is height on the shore, which determines the ratio of emersion to submersion. Both algal growth rates and grazing pressure also differ in
pools and on open rocks.
An important point here is that growth rates and grazing rates tend to be influenced by the same factors, sometimes in the same direction, sometimes in opposite
directions. These factors are all concerned with grazer effects on single algal species. Community effects of grazing depend on the relationship between grazer
preferences and algal competitive hierarchies. These hierarchies may differ between habitats, particularly open rocks and pools.
I shall emphasize the importance of height on the shore by considering the ecological effects of species found at different heights.
High shore—Nodilittorina unifasciata

I follow Reid (1989) in using the name Nodilittorina unifasciata for the animal referred to throughout the ecological literature as Littorina unifasciata. Nodilittorina
unifasciata is one of three littorines that characterize the upper shore of New South Wales (Underwood 1981), although it also occurs on the mid shore (Jernakoff
1986). It eats epilithic microalgae in preference to macroalgae (Jernakoff 1983, Black et al. 1988), but Branch & Branch (1981) found that highshore populations
feed mainly on lichens.
Fairweather et al. (1984) highlight the fact that relations between predators and motile prey such as littorinids are complex and variable. The effects of predation on
prey populations can vary enormously over small distances (tens or hundreds of metres) on rocky shores (Fairweather & Underwood 1991) and there is also
considerable variability in the effects of predators (mostly whelks or crabs) on N. unifasciata populations, depending on the geographical area and height on the shore.
This is caused by many interacting factors, including not just the density of predators, but also the prey preferences of predators under different circumstances.
Individual whelks can have much narrower diets than field observations of feeding by the species suggest. West (1986) found that, although Nucella emarginata took
seven species of prey in the field, no individual took more than three species in four months. Marked differences in growth rate among conspecific whelks may be due
to differences in diet (Moore 1938, Moran et al. 1984). The whelk Lepsiella vinosa feeds primarily on mussels in zones where mussels and littorinids are available.
Individuals eating mussels both grow
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faster and produce more eggs than those that live higher on the shore, where mussels do not occur, and feed on Nodilittorina unifasciata (Ward & Quinn 1988). This
may be partly explained by the effect of submersion period on rate of feeding, but these authors attribute the preference for mussels, where both prey are available, to a
lower energetic profitability for N. unifasciata. Field estimates may badly underestimate rates of N. unifasciata predation by whelks because of their short handling
time (Fairweather & Underwood 1983), but highshore littorinids seem rarely to be eaten where alternative prey are present (Fairweather 1988, Ward & Quinn
1988). They can, however, form the bulk of whelk diets where they are the most abundant prey (McQuaid 1985, Ward & Quinn 1988).
Branch & Branch (1981) assume that predation is not an important regulator of N. unifasciata populations, but there is equivocal evidence that the presence of
whelks may influence local densities of N. unifasciata. Jernakoff & Fairweather (1985) found that the presence of Morula marginalba did not affect the rate of
recolonization of scraped areas by Nodilittoria unifasciata, but Fairweather et al. (1984) found that densities of N. unifasciata rose where Morula was removed and
declined in areas with whelks. They conclude that this difference could be explained by predation, immigration or recruitment of small snails. Fairweather &
Underwood (1991) attributed a decrease in Nodilittorina unifasciata numbers where Morula was present to snails fleeing, although some were eaten. Preferential
immigration into areas without whelks could be consistent with the predator avoidance behaviour shown by Nodilittorina unifasciata. McKillup (1981) found that N.
unifasciata showed chemosensorybased avoidance of whelks in the field. He demonstrated the selective advantage of this by exposing New Zealand populations of
N. unifasciata to an Australian whelk, Lepsiella vinosa, of which they had no experience. Although Australian Nodilittorina unifasciata, which normally cooccur
with Lepsiella vinosa, avoided the whelk, New Zealand Nodilittorina unifasciata did not and were eaten. Whatever the mechanism for reducing N. unifasciata
densities in the presence of whelks, it is possible that this will produce local variations in littorinid grazing intensity.
Apart from possible effects of predators, the local density of N. unifasciata may also be influenced by the availability of suitable shelter. Underwood & Chapman
(1985, 1989) found that topographic complexity, in the form of crevices, depressions, etc., affected directionality and rate of movement of N. unifasciata. Unlike N.
pyramidalis (Chapman & Underwood 1994), topography did not affect local densities of N. unifasciata. Nor is there any indication that on the high shore, where
foliose macroalgae are absent, local variations in density affect the rest of the community (Underwood & Chapman 1992).
Branch & Branch (1981) conducted caging experiments on highshore N. unifasciata. They showed a reduction in body weight with increased caging densities and
suggested that natural populations suffer intense intraspecific competition. They also found that snails dispersed away from areas of low food availability, but tended to
remain where there was more food. In Western Australia N. rugosa occurs immediately downshore of N. unifasciata (Black et al. 1979) and Branch & Branch refer
to unpublished data from Ayre et al. showing that removal of either N. unifasciata or N. rugosa allows the other species to expand its vertical range.
Lower down the shore, Jernakoff (1986) found that barnacles increase local densities of N. unifasciata. Chapman (1994), however, found that a positive
correlation between the densities of barnacles and N. unifasciata only held at some sites. Such a correlation may affect other components of the community as foliose
macroalgae grow in this zone. The effects of grazing by midshore populations of N. unifasciata have been studied experimentally by Jernakoff (1983). Because N.
unifasciata eats macroalgae only if microalgae are unavailable,
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sporelings that avoid initial grazing will be safe from future grazing as long as other microalgae are present. In this way the distribution of adult macroalgae reflects the
history of sporelings that “escaped” grazing. In contrast to limpet grazing in Oregon (Cubit 1984), as well as North American and European populations of Littorina
littorea (see below), grazing pressure by Nodilittorina unifasciata does not vary with season. Nevertheless, the effects of grazing may vary seasonally because of the
interplay of grazing with algal growth rates. For example, Williams (1993) found that grazing by Nodilittorina spp. decreased in summer on the midshore in Hong
Kong as grazers moved farther downshore, In this case the high temperatures of summer were more important, as they reduced algal abundance dramatically and
masked the effects of herbivores.
Mid shore—Bembicium nanum

The mid shore in New South Wales is dominated by barnacles at exposed sites and high densities of grazers on sheltered shores. With intermediate exposure there is a
mixture of barnacles and grazers (Underwood et al. 1983). Bembicium nanum forms about 12% by number of the grazers on these mid shores (Underwood &
Jernakoff 1984). It is an epilithic grazer and the only member of the genus found on rocky shores (Reid 1988). B. nanum feeds on microalgae (including sporelings)
and on ephemeral macroalgae such as Ulva and Enteromorpha (Underwood 1984b, Quinn & Ryan 1989). It is more abundant on waveexposed shores and where
there are fewer pools (O’Gower & Meyer 1971, Underwood 1975). Lowtide distribution is clumped (O’Gower & Meyer 1971, Underwood 1976a) but, although
the abundance of Bembicium nanum increases with greater cover of the encrusting alga Peyssonelia gunniana, this is not due to habitat preferences. The distribution
of Bembicium nanum is random with respect to the presence of Peyssonelia (Underwood 1976b).
In New South Wales littorinids are only occasionally eaten by the whelk Morula marginalba (Fairweather 1988), and Bembicium nanum showed no decrease in
survival where whelks were present compared with areas of open rock where they were removed (Fairweather et al. 1984). Fairweather & Underwood (1991),
however, report an increase in numbers of B. nanum in pools from which whelks were removed. Morula in these pools selected smaller individuals and Bembicium
could grow into a refuge in size within two years. In South Australia crabs are abundant and prey extensively on B. nanum. By selecting particular size classes they may
be responsible for size gradients up the shore (Chilton & Bull 1984, 1986).
Grazers have important effects on macroalgae on these shores, but their effectiveness in controlling algal cover and species composition depends partially on the
interaction of grazing with factors that influence algal growth rates. These include height on the shore and degree of wave action (e.g. Underwood & Jernakoff 1984).
Grazing is the major determinant of the upper limits of distribution for many species of algae on these shores (Underwood & Jernakoff 1984). Underwood (1980) and
Underwood & Jernakoff (1984) have shown that some macroalgae can survive higher on the shore if grazers are excluded. Macroalgae may, however, suffer from
physical stress higher upshore, unless sheltered by cages, and form turfs rather than full sized plants. Where turfs are allowed to develop and then exposed to
herbivores, they are removed by direct invasion by macroalgal grazers such as the limpet Siphonaria, while microalgal grazers such as Bembicium nanum remove
turfs from the edge inwards (Underwood 1980).
Greater wave action tends to increase colonization and growth of algae. While foliose macroalgae are abundant on the low shore, they are rare or absent on the mid
shore of shel
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tered areas (Underwood 1981, 1984c, Underwood & Jernakoff 1981, 1984). On sheltered shores grazing eliminates the sporelings of most foliose and ephemeral
algae, leaving only perennial crustose forms. Most crustose cover is formed by Hildenbrandia prototypus that is much less susceptible to grazing than foliose species.
Herbivores feeding on its surface mainly remove ephemeral algae. An important point is that H. prototypus is able to spread from small pits or crevices that are
inaccessible to grazers. Because of this it does not need a large number of ungrazed sporelings to occupy a lot of space (Underwood 1980).
As with Littorina littorea in New England, microalgal grazers remove ephemeral algae, but in this case there is no evidence that ephemerals have a marked effect
on the dominant perennial species. Thus Bembicium nanum and the other grazers have no longterm effect on interactions among the algae in New South Wales
(Underwood 1980). However, Braley et al. (1991) suggest that, in Victoria, B. nanum does influence algal interactions. They found a negative correlation between the
densities of B. nanum and the most abundant perennial macroalga, Hormosira banksii. Bembicium nanum did not significantly affect recolonization of cleared areas
by Hormosira banksii, but did reduce recolonization by the ephemeral alga Scytosiphon lomentaria that overgrew at least one patch of Hormosira banksii
sporelings. Although the experiment was very short (eight weeks) the results suggest that removal of Scytosiphon lomentaria by Bembicium nanum is beneficial to
Hormosira banksii. This does not accord with the negative correlation between H. banksii and Bembicium nanum.
The abundance of microalgae on shores in New South Wales increases downshore except in summer when it decreases abruptly at all levels (Underwood 1984c).
There is, however, no correlation between these vertical or seasonal changes and herbivore abundance and Underwood (1984b) found that B. nanum could grow
when caged at four heights across the shore, although growth was slower at the top level. Growth rate was not correlated with food availability or the duration of
submersion (or the product of the two) and he suggests that food availability may have been sufficient to support maximum growth rates at all except the top level. The
neritid Nerita atramentosa only grew at the two lowest heights although Bembicium nanum was competitively inferior to both it and the limpet Cellana tramoserica
in caging experiments (Underwood 1978). All three showed greater weight loss and mortality as total density increased, but the presence of Bembicium nanum did
not affect either body weight or mortality of the other two. These results were ascribed to the feeding biology of the species as B. nanum removes fewer microalgal
cells from the substratum than the other two species (Underwood 1984a). Cellana tramoserica has a higher rate of radular action in the field (Petraitis 1992) and
excavates more deeply than the others (Underwood 1984a), while Nerita atramentosa moves further and feeds faster than Bembicium nanum, which moves
randomly over short distances when feeding (Underwood 1977, 1978). Petraitis (1992) suggests that the greater sensitivity of B. nanum to disturbance while grazing
may influence the consistency of these competitive interactions. Cellana tramoserica had the strongest effect on Bembicium nanum, but under natural conditions
would suffer from intraspecific effects before reaching densities high enough to affect B. nanum. Negative interaction between Nerita atramentosa and Bembicium
nanum could happen in the field and the two species do clump separately at low tide, although this is ascribed to nonfeeding effects as they both graze at high tide
(Underwood 1976b).
Caging experiments in Victoria involving the trochid Austrocochlea constricta and Bembicium nanum support several of these findings. This population of B.
nanum was not food limited, despite lower food availability and body weights in summer (Quinn & Ryan 1989). B. nanum experienced no significant inter or
intraspecific competitive effects (weight loss or mortality) even at artificially high densities. Austrocochlea constricta feeds
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on diatoms that can be overgrown by macroalgae, and Quinn & Ryan suggest that Bembicium nanum may facilitate feeding by the trochid by removing macroalgal
sporelings and ephemeral algae.
Low shore—Littorina littorea

Littorina littorea is an example of an epilithic grazer found on open rocks on the low and mid shore as well as in pools in New England and Europe (Ballantine 1961,
Lubchenco & Menge 1978, Lubchenco 1983). There has been particular interest in this species in North America where it has become invasive and has had a
profound influence on community structure (Carlton 1992, Vadas & Elner 1992). Its effect on algal communities in Europe has been less well studied. On European
shores L. littorea is generally much less abundant and ecological research has focused more on limpets (Norton et al. 1990). A major exception is the work of Janke
(1990) that must be considered before the work in North America.
Janke examined interactions of herbivory, predation and competition for space at different heights on a sheltered shore on Helgoland in the North Sea. Of the three
species of Littorina he investigated, only L. littorea had a marked effect on algal populations, and this was only on the mid shore. On the high shore abiotic factors
(especially wave action) had an overriding effect and grazers (L. saxatilis and L. littorea) at natural densities had little influence on algal abundance. On the low shore,
algal growth was only slowed by grazing if L. littorea density was doubled. On the mid shore, however, natural densities of L. littorea excluded ephemeral algae
(Ulva and Enteromorpha) and reduced the cover of Fucus spp. This has ramifications as macroalgae and mussels compete for space in this zone.
In a series of short term experiments in the Wadden Sea, Wilhelmsen & Reise (1994) found that grazing control of ephemeral algae by Littorina littorea was
strongly density dependent. L. littorea prevented the development of macroalgal beds on mussels, where the snails occurred at high densities, but not on soft sediment,
where snail density was low.
Historical background to L. littorea in North America There is controversy over whether L. littorea is exotic or indigenous to North America. Clarke (1971)
suggests that it is both! Records of L. littorea in North America before the mid nineteenth century are rare (Vermeij 1982), but there is evidence for its occurrence well
before the last century (Clarke & Erskine 1961, Clarke 1963, Bird 1968, Wagner 1977). However, even these early populations are assumed to have originated in
Europe. Kraeuter (1974) calculates that the time required for plankton to cross the Atlantic is too long for direct colonization by larvae and suggests they were
introduced via Iceland and Greenland. He postulates the establishment of steppingstone populations there during the postglacial climate optimum (5000–3000 BC)
and their subsequent elimination by falling temperatures and expansion of the Arctic packice around 900–450 BC. Subfossils of L. littorea have been found in both
Spitsbergen (Salvigsen et al. 1992) and Iceland (references in Kraeuter 1974), although it does not occur in either area now.
Earlier North American populations seem to have been restricted to eastern Canada (Vermeij 1982), and the sudden and rapid range expansion of L. littorea
beginning in the mid nineteenth century has been explained by recolonization as shipping between the new and old worlds increased. Clarke (1971) suggests that early
populations were restricted by low physiological plasticity and that shipping caused an infusion of new genetic material that allowed the species to take advantage of a
general rise in sea temperatures around 1860–70. The subsequent spread of L. littorea southwards is well documented (see references in
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Brenchley & Carlton 1983) and it rapidly became one of the most abundant snails in the intertidal and shallow subtidal of the northeastern United States.
Community effects The vertical and horizontal distribution of the major species on rocky shores of New England are illustrated by Vadas & Elner (1992). Population
expansion of L. littorea in New England may have been possible partially because of the absence of predators and partially because of the absence of competitors.
Menge & Sutherland (1976) compare the east and west coast intertidal systems of the northern USA. Herbivore guilds are smaller on the east coast and there is only
one midshore invertebrate predator. Crabs and starfish are confined to the low shore and subtidal while fish and birds feed infrequently on the species in this food
web. L. littorea in New England is preyed on by the green crab Carcinus maenas (Hadlock 1980), which is also exotic to New England. Littorina littorea shows a
strong alarm response to predation by Carcinus maenas, but in the Bay of Fundy it is only taken by larger crabs and even these prefer Littorina obtusata in the
laboratory (Rangeley & Thomas 1987). Unlike individuals preyed on by Cancer pagurus in Europe (Lawton & Hughes 1985), Littorina littorea has a refuge in size
from Carcinus maenas (Hylleberg & Christensen 1978). There is no indication that Littorina littorea populations are controlled by predation and they reach very
high densities in some habitats.
Not only is L. littorea abundant on the eastern coast of North America, it also has a profound influence on community structure. L. littorea has important effects on
sessile organisms, other grazers and even the existence of certain habitats. The effect on community structure depends partly on its abundance. For example, at high
densities L. littorea can clear areas of the ephemeral alga Enteromorpha, while at lower densities it can maintain, but cannot establish, cleared areas (Petraitis 1983).
Density in turn depends largely on the degree of wave exposure. Density varies considerably with site and habitat (Table 1), but generally Littorina littorea is most
abundant, and has its greatest effects, on sheltered shores and in pools. It becomes more rare as wave action increases and is excluded entirely from very exposed
shores. This is true in both Great Britain (Moore 1940, Ballantine 1961) and New England (Lubchenco & Menge 1978). The abundance and activity of L. littorea are
also both patchy and seasonal (Janke 1990, Vadas 1992). Littorinids were much less affected than top shells (Monodonta lineata and Gibbula spp.) during an
exceptionally cold winter in Europe in 1963 and Littorina littorea showed no signs of increased mortality (Crisp 1964). Nevertheless, its rate of feeding on fucoids at
5°C is about half that at 15°C (Barker & Chapman 1990) and in some places L. littorea migrates downshore during winter to the low intertidal (Bertness 1984) or
subtidal (Lein 1980).
Effects on algae The results of interactions among L. littorea and algae are very complex for several reasons, (a) L. littorea affects different algae in different ways;
(b) it affects different life history stages of the same alga in different ways; (c) it may affect the same stage of the same alga differently under different physical conditions
or in the presence of other algae; d) the algae interact among each other.
L. littorea is an omnivorous grazer, it incidentally eats sessile animals or egg masses (see below), but feeds primarily on microalgae (including sporelings) and
macroalgae (Newell 1958, Hylleberg & Christensen 1977, Lubchenco 1978, Brenchley & Carlton 1983, Petraitis 1983). Davies et al. (1992) suggested that mucous
trails might be important as a food source, but found that they are relatively shortlived and inefficient at trapping diatoms. Grazing movements of the head and the
pattern of bite marks made by the radula are described in Juch & Boekschoten (1979). L. littorea has a clear order of species preferences among
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Table 1 Densities of Littorina littorea reported in the literature. Densities can be extremely patchy; e.g. Vadas (1992) recorded mean densities of
250m−2 on the low shore in spring at a site in Maine, but densities in small algal patches were up to 4000m−2.
Degree of exposure Density (no. m−2) Location Source

North America
Exposed 0 Nova Scotia Barker & Chapman 1990
Semiexposed 2 Massachusetts/Maine Lubchenco 1982
Sheltered (mid shore) 274 Massachusetts/Maine Lubchenco 1980
Sheltered (low shore) 650 Massachusetts/Maine Lubchenco 1980
Sheltered bay 400 New York Petraitis 1983
Sheltered bay 100–650 Gulf of Maine Petraitis 1987
Sheltered bay 400–800 Rhode Island Bertness 1984; Bertness et al. 1983
Exposed pools 0–0.4 Maine Sze 1982
Exposed pools 200–1200 Nova Scotia Barker & Chapman 1990
Semiexposed pools 14–38 Maine Sze 1982
Semiexposed pools 2811 Massachusetts/Maine Lubchenco 1982
Sheltered pools 4276 Massachusetts/Maine Lubchenco 1982
Sheltered pools 120–457 Maine Sze 1982
Spartina beds 124–360 Massachusetts Brenchley & Carlton 1983
Spartina beds 184 Massachusetts Brenchley 1982
Europe
Extremely sheltered >50 South Wales Ballantine 1961
Very sheltered 10–50 South Wales Ballantine 1961
Very sheltered 40–100 Helgoland, Germany Janke 1990
Sheltered 15–260 Norway Lein 1980
Sheltered 100–350 Wadden Sea Wilhemsen & Reise 1994
Moderately exposed <1−>250 Isle of Man Norton et al. 1990
macroalgae, favouring ephemeral species such as Enteromorpha intestinalis above brown algae such as Fucus vesiculosus, while Ascophyllum nodosum, the red
alga Chondrus crispus and encrusting algae are not eaten at all (Lubchenco 1978, 1980, 1982, Bertness et al. 1983, Watson & Norton 1985a). Feeding preferences
are influenced by macroalgal defences, both chemical and mechanical, which are considered below in the section on macroalgal grazers (pp. 285–6).
A preference for ephemeral algae is reflected in a negative correlation between Enteromorpha intestinalis abundance and density of Littorina littorea that has
been recorded both in space (Sze 1980, Petraitis 1983) and in time (Lein 1980). Lein (1984) suggests that feeding preferences of L. littorea may even be partially
responsible for the restriction of Fucus ceranoides to estuarine areas in Norway, despite its ability to tolerate high salinities. In areas with particularly dense populations
of Littorina littorea, castup seaweed flotsam (wrack) may be an important source of food. L. littorea feeding on wrack consisting of mixed species, show the same
preferences as with attached algae (Watson & Norton 1985a).
On sheltered rocky shores L. littorea generally decreases algal diversity and the abundance of preferred food species. It also mediates interactions between
competing algae through feeding preferences. Grazing by L. littorea removes ephemeral algae so that algal diversity decreases as snail density increases (Lubchenco
1978). This tends to speed up succession (Lubchenco 1983), but it also influences the perennial algal species directly. Fucus spp. can survive well on the low shore in
New England, but under natural conditions are
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competitively excluded by Chondrus crispus. Where C. crispus is experimentally removed the abundance of Fucus spp. depends on the intensity of Littorina
littorea grazing (Lubchenco 1980), Higher up, on the mid shore, removal of ephemerals by grazing releases Fucus vesiculosus from competitive inhibition and allows
it to dominate space more rapidly. F. vesiculosus sporelings are susceptible to grazing but can, if they escape grazing, grow into a refuge in size that is correlated with
higher concentrations of defensive chemicals in larger plants (Denton et al. 1990). Grazing actually benefits larger fucoids by removing epiphytes (Lubchenco 1983).
Lubchenco suggests that F. vesiculosus sporelings can escape grazers by three means. Support for each of these suggestions comes from the work of other authors.
1. Sporelings may survive in microscale refuges such as crevices or among barnacles where grazing is reduced. This is supported by the work of others on Littorina
littorea (Menge 1976, Bertness et al. 1983, Petraitis, 1987), but not by Jernakoff (1983) working on an assemblage of grazers including Nodilittorina
unifasciata. These different findings may reflect differences in the body sizes of the grazers (Geller 1991, see below).
2. Grazers may simply miss sporelings due to patchiness in grazing (e.g. Jernakoff 1983). Petraitis (1983) showed that Littorina littorea can consistently miss areas
because of nonrandom grazing patterns. Directional (i.e. nonrandom) movement is caused by disturbance so this effect may be greatest where snails are disturbed
more frequently by wave dislodgement (Petraitis 1982). Thus L. littorea may not only be less abundant on more exposed shores, it may also graze the shore less
efficiently.
3. Herbivores may be swamped by high sporeling densities and growth rates (e.g. Underwood & Jernakoff 1981).
Other workers have recorded the devastating effect of L. littorea grazing on sporeling survival (e.g. Barker & Chapman 1990). Miller & Vadas (1984) report that
grazing by L. littorea has a marked effect on survival of Ascophyllum nodosum germlings, even though mature plants contain polyphenols, and Watson & Norton
(1985a) showed that snails dislodged A. nodosum germlings, even when not feeding on them. Watson & Norton also found that species preferences of Littorina
littorea for germlings mirrored those for mature plants, ephemerals being preferred over perennials. However, while adult Ulva lactuca was preferred to germlings,
the opposite was true for Fucus serratus. They attribute this difference to the relative efficiency with which mature plants and sporelings of the same species may be
eaten.
Central to our understanding of grazing effects is the point highlighted by Vadas & Elner (1992); grazing effects can be very patchy. Local densities of Littorina
littorea can vary over short distances because of slight variations in wave action and snails can form destructive feeding fronts at critical densities. Vadas & Elner
interpret the main effect of L. littorea as the recycling of free space by removing ephemeral algae from disturbed patches. Patches of seasonal ephemeral algae within
stands of perennial algae can show longterm persistence, the same patches being recycled through the activities of L. littorea and the whelk Nucella lapillus for at
least 17 years (Vadas 1992). Janke (1990) also observed very high densities of Littorina littorea in small patches where ice scour removed macroalgae and noted
that these patches remained open for over two years.
Littorinids can be important grazers in tidal pools. Trowbridge (1993) estimates that, during summer, L. scutulata accounts for over 90% of grazing on the
filamentous alga Chaetomorpha in highintertidal pools in Oregon. The effect of Littorina littorea on algal interactions differs between open rocks and pools. In
pools, grazing intensity is more patchy and algal competitive hierarchies may be reversed. Pools may contain no littorinids, or they
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may support very high densities. Snails in pools may also exert greater grazing pressure as they remain submerged and can feed throughout the tidal cycle.
Lubchenco (1978) examined densitydependent grazing effects of L. littorea in highshore pools. As the preferred food species (Enteromorpha intestinalis) was
competitively dominant in sheltered areas, highest algal diversity occurred at intermediate snail densities. Intense grazing removed most species while low snail density
allowed E. intestinalis to dominate. In a nonexperimental study, Sze (1980, 1982) also found that Littorina littorea was associated with mixedspecies algal
communities in New England tidal pools and that algal species richness was lower at high densities of snails. However, no algae were positively associated with high
densities of snail. This contrasts with Nova Scotia where pools dominated by Fucus distichus are associated with very high densities of Littorina littorea (Barker &
Chapman 1990).
Again wave exposure is important, because it influences competitive hierarchies. Sze (1980) found that, where L. littorea was absent, algal communities were mixed
at exposed sites, but Enteromorpha intestinalis dominated in shelter. An interesting consequence of this, described by Lubchenco (1978), was that, once well
established, E. intestinalis formed a canopy that enhanced survival of the crab Carcinus maenas. C. maenas eats small Littorina littorea producing conditions for
persistent low snail density in pools dominated by Enteromorpha intestinalis and crabs. In midshore pools at sheltered sites in New England, herbivores (primarily
Littorina littorea) were partially responsible for the exclusion of Fucus vesiculosus (Lubchenco 1982). Sporelings are less likely to escape grazing in pools because
the abundance of snails can be much higher than on open rocks. In sheltered bays grazing by snails is intense enough to exclude F. vesiculosus from the open rock as
well as pools and even plants grown for a year in cages only occasionally survived subsequent exposure to natural grazing rates (Petraitis 1987).
These studies clearly portray Littorina littorea as a grazer with profound effects on community structure. But the application of this perception to the northwestern
Atlantic as a whole has recently been challenged (Barker & Chapman 1990, McCook & Chapman 1991, 1993). Chapman and his coworkers found that, although L.
littorea was the only littorinid grazer in Nova Scotia that had a significant effect on sporeling survival and thus on algal interactions, it was largely restricted to pools.
They also found that grazing intensity depended not only on the species of alga considered, but also on what other species were available. Of four species of Fucus, F.
vesiculosus was preferred by Littorina littorea (and L. obtusata) in the laboratory, but in the field Fucus evanescens was more heavily grazed. Their experimental
design allowed access to many different grazers including amphipods and isopods. Clearly where Littorina littorea is less abundant, as on exposed shores in Nova
Scotia, its effect on community structure is outweighed by other grazers.
L. littorea also has community level effects on microalgae. Hunter & RussellHunter (1983) examined the effects of grazing by L. littorea on microalgae grown on
glass slides. These microalgae formed a high diversity assemblage, predominantly diatoms, existing in three dimensions. There was no active selection of diatoms, but
there was a degree of passive selection in feeding as some diatoms were less likely to be removed by grazing, for example small species cemented to the substratum.
Because there were no competitive dominants, and no active selection during grazing, the effects were akin to those on macroalgae on open rocks, rather than in pools
(Lubchenco 1978). Grazing resulted in low abundance, low diversity, twodimensional assemblages. As grazing intensity increased, diversity decreased. Moderate
grazing also caused an increase in productivity and the C:N ratio of the overall community increased with grazer density.
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Effects on sessile and sedentary animals Interactions among littorinids, barnacles and algae are complex because each of these elements can influence the other. For
example, not only do barnacles affect littorinid grazing, but littorinid grazing can affect barnacle recruitment. Small or juvenile littorinids often shelter between barnacles
or in the tests of dead barnacles (Moore & Sproston 1940, Hawkins 1981, Underwood & McFadyen 1983, Geller 1991). Jernakoff (1986) found that the quality of
barnacles as shelter depends on algal cover. In New South Wales, barnacle tests without algal cover were rapidly washed away, while those with a complete cover of
algae remained on the shore, but were not used by littorinids as they accumulated sediment that became anoxic. Studies on large littorinids, like L. littorea, have found
that barnacles can interfere with littorinid grazing (Bertness et al. 1983, Lubchenco 1983). Alternatively, high densities of small littorinids, such as L. neglecta, sheltering
among barnacles can produce local areas of high grazing intensity and low algal biomass (Hawkins 1981). The difference in these findings depends on the size of the
littorinid. Geller (1991), working in California, showed that the effect of epilithic grazers on algal colonization depends on body size and can be masked by the effects of
other grazers. By removing limpets and/or L. plena he showed that L. plena alone cannot control the development of algal cover on open rocks, but that it is the more
effective grazer among barnacles. He also found that small L. plena had a stronger inhibiting effect on algae than large individuals.
Menge (1976) found that L. littorea had no effect on barnacle or mussel recruitment at natural densities. But yet again the degree of exposure is important. In
sheltered bays L. littorea affects sedentary animals both directly and indirectly (Petraitis 1983, 1987). Intermediate levels of grazing by L. littorea enhance barnacle
recruitment. At low snail densities barnacles are fouled by ephemeral algae and sediments, while at high densities snails can directly graze newly settled cyprid larvae.
Again, this is made more complex because barnacles can influence the distribution of grazers and thus algal cover and the success of barnacle recruitment (Jernakoff
1985, Underwood et al. 1983).
The effects of L. littorea on mussel recruitment are less direct and less predictable. Petraitis (1987) found that mussels recruited directly onto barnacles and so
benefited from greater barnacle densities, which are in turn affected by the activities of L. littorea. Mussels often, but not always, settle initially on filamentous algae
(Bayne 1964, McGrath & King 1991) and Petraitis (1990) suggests that L. littorea may also depress mussel recruitment by removing algae. Petraitis (1987) lists a
number of other confounding effects that are potential consequences of littorinid grazing and that produce a network of interacting factors. For example, with less
extreme grazing macroalgae can not only reduce barnacle recruitment, but also enhance the effectiveness of mussel predators by offering them shade.
Effects on other grazers Such an abundant grazer as L. littorea may be expected to have a marked effect on other herbivores, but this is not always the case. In
England, L. littorea and the gastropod Hydrobia ulvae coexist in saltmarshes, although the two species tend to occur on different substrata. Removal of Littorina
littorea from experimental plots for 3month periods had no effect on Hydrobia densities (Frid & James 1988). Littorina littorea and L. obtusata cooccur on hard
substrata, but are specialized to feed on different algae and occupy different microhabitats (Watson & Norton 1985a,b, Barker & Chapman 1990). The presence of L.
littorea has no effect on survival, growth or habitat selection by L. obtusata (Seeley 1982).
On the other hand, Vermeij (1978) correlates a contraction of range of the saltmarsh snail Littoraria irrorata in New England with southward expansion of
Littorina littorea. The
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impact of L. littorea has also been assessed for three other herbivores, all showed that it had a negative influence, but to varying degrees.
Field experiments have been used in New England to examine interactions between Littorina littorea and both L. saxatilis (Behrens Yamada & Mansour 1987)
and the limpet Notoacmea testudinalis (Petraitis 1989). The experiments were of different designs and durations, but both showed inhibition of growth when
experimental animals were caged with Littorina littorea. L. saxatilis generally lives higher on the shore, but cooccurs with L. littorea on the upper shore at Woods
Hole in Massachusetts. Both species grew faster when caged lower on the shore, as do the neritid Nerita atramentosa (Underwood 1984b) and the trochid Oxystele
variegata (McQuaid 1982), but on the mid shore Littorina saxatilis very rapidly (18–19 days) showed significant depression of growth when caged with L. littorea.
L. littorea had higher growth rates than L. saxatilis and grew faster when caged with L. saxatilis than with conspecifics. Behrens Yamada & Mansour (1987) suggest
that the upper limits of distribution of L. littorea on the shore are set by food availability.
Petraitis (1989) ran longer experiments (59–77 days), which showed that caging with L. littorea resulted in decreased growth and survival of Notoacmea
testudinalis. There was no significant difference between mean body weight of limpets caged with 25 or 50 littorinids in experimental baskets, but Petraitis suggests
that the presence of a few littorinids may benefit limpets by removing filamentous algae, without affecting the microalgae on which the limpets feed (see also Quinn &
Ryan (1989) above on Bembicium nanum). In neither of these studies do the results indicate whether the presence of Littorina littorea influences distribution or
abundance of the other herbivore.
The third case provides striking evidence that L. littorea does exactly that. L. littorea occurs at high densities in very sheltered areas such as saltmarshes, where it
occurs mainly on hard substrata (plant stems, etc.). Frid & James (1988) concluded that competition between L. littorea and Hydrobia ulvae in an English saltmarsh
was minimized because of only partial overlap of diet and habitat use. Dippolito et al. (1975) used similar reasoning to claim that, in New England, Littorina littorea
was not displacing the indigenous mud snail Ilyanassa (Nassarius) obsoleta. Later work by Brenchley (1980, 1982) and Brenchley & Carlton (1983) refutes this.
Physical contact with Littorina littorea is actively avoided by Ilyanassa obsoleta, which vacates pools to which littorines are added. If Littorina littorea is removed
from pools Ilyanasa obsoleta density is not affected, but it extends its microdistribution onto the hard surfaces previously occupied by Littorina littorea. This is
important because Ilyanassa obsoleta lays egg masses on hard substrata and contact with L. littorea delays egg laying. Although Littorina littorea does not actively
seek out egg masses, it ingests them incidentally while feeding and is so abundant that it is one of the two major predators on egg masses (Brenchley 1982).
Avoidance behaviour affects the entire system because it influences the migration of Ilyanassa obsoleta down into the subtidal during winter and upshore in the
spring. Migration takes it through lowshore beds of Zostera, but if these are occupied by Littorina littorea they can act as barriers. Mud snails trapped upshore of
such barriers hibernate in the midshore sediments instead of migrating. Other animals returning from the subtidal in spring are channelled through gaps between Zostera
beds on their way upshore. Rather than dispersing over the shore as usual at densities of around 500m−1, these animals form dense swarms of well over 1000m−1.
These swarms retain their cohesion for the summer and result in quite different bioturbation effects on the sediments from normal. Low densities of Ilyanassa obsoleta
stimulate algal growth in the sediments, while high densities have negative effects. Infaunal distribution and abundance correlate with density of mud snails and are most
patchy where
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mud snails form swarms (Brenchley 1980, Connor et al. 1982). There is evidence from England that Littorina littorea itself also affects the distribution of the infauna
of saltmarsh sediments (Frid & James 1988).
Ilyanassa obsoleta is also excluded from marsh grass beds at the top of the intertidal and extends farther upshore if Littorina littorea is experimentally removed.
Despite this evidence for habitat displacement of Ilyanassa obsoleta and its exclusion from some parts of the shore, Brenchley & Carlton (1983) conclude that
littorinids do not affect the geographical distribution of the mud snail as it has refuge habitats (mud) in which Littorina littorea is rare. Another study, however, suggests
that the presence of L. littorea may threaten the very existence of this habitat (see below).
Effects on nongrazing motile animals For completeness I shall mention a final effect of L. littorea. Scully (1979) examined shell use by the hermit crab Pagurus
longicarpus at Rhode island. At both of his sites Littorina littorea was the most abundant gastropod and supplied most of the shells used. Where hermit crabs
depend on indigenous shells, changes in their populations can be highly correlated with the availability of shells (e.g. Spight 1977). However, L. littorea densities are so
high that Scully (1979) suggests that Pagurus longicarpus is not limited by the availability of shells, even though this is highly seasonal. Consequently, P. longicarpus
may be more numerous than before the spread of Littorina littorea (Carlton 1982, in Brenchley 1982). Brenchley (1982) suggests that this may increase pressure on
the mud snail Ilyanassa obsoleta as, like Littorina littorea, hermit crabs are important predators of its benthic egg cases. Increases in hermit crab populations may
also affect the abundance of symbionts and shell epifauna (which can be associated with particular shell/crab species combinations) via the “habitat web” described by
McLean (1983).
As well as population density, the availability of L. littorea shells may affect the size structure and sex ratios of populations of Pagurus longicarpus. Hermit crabs in
a saltmarsh in Florida (beyond the range of Littorina littorea) have an ample supply of Littoraria irrorata shells, but these are not large enough for the bigger crabs.
Large crabs may be shell limited in spite of the abundance of small shells (Wilber & Herrnkind 1984). In an area where Littorina littorea does occur (Connecticut),
small hermit crabs mostly use indigenous gastropods shells, but large males use the shells of L. littorea (Blackstone & Joslyn 1984). This represents an ontogenetic
shift in shell preference of the crabs rather than shell availability and mainly affects males as females do not grow as large. Very few indigenous shells of comparable size
were found (mostly Ilyanassa obsoleta and Urosalpinx cinerea) and the increase in availability of large shells must release large crabs from shell limitation.
Habitat modifications Bertness (1984) found that the activities of Littorina littorea in sheltered bays resulted in a change in habitat type. Experimental removal of
littorinids resulted in rapid sedimentation of rock surfaces. Snails bulldozed sediments from the rock surface and when they were excluded, sedimentation was further
increased by the establishment of an algal canopy. L. littorea alters the habitat from one favouring sedimentdwelling organisms such as polychaetes, to one favouring
hardsubstratum species such as crustose algae and barnacles. L. littorea also has a significant effect on the marsh grass Spartina alterniflora. As well as reducing the
accumulation of sediments in which S. alterniflora roots, Littorina littorea eats both shoots and rhizomes. Removal of snails led to an expansion of Spartina beds, as
well as an increase in their productivity. In view of its recent arrival, or at least range and population expansion, in North America, Bertness concludes that intertidal soft
sediment habitats and fringing saltmarshes may have been more common in the past than at present.
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Historical perspective on species interactions Vermeij (1978) describes the rocky shore biota of New England as an impoverished version of the North European
biota (see also Norton et al. 1990). He suggests that this is linked to past glaciation events and the absence, between New York and Florida, of rocky shores that
could form refuge habitats for species forced south by glaciation. Patellid limpets and top shells are absent and have no ecological equivalents while one of the most
important grazers, Littorina littorea, is exotic, or at least invasive. The fact that both fucoids (Geiselman & McConnell 1981, Barker & Chapman 1990) and crustose
algae (Bertness et al. 1983) in New England have chemical defences that deter L. littorea suggests that the phylogenetic lines of these algae evolved in environments
with naturally intense grazing regimes.
The situation resembles that described by Berman & Carlton (1991) for an invasive pulmonate that cooccurs with Assiminea californica and Littorina
subrotundata in Oregon saltmarshes. Rather than having displaced indigenous species through competition, L. littorea has entered an empty niche, unoccupied by
ecologically equivalent local species. Not only is L. littorea less abundant in Europe (see Table 1 p. 273), it also has a more restricted vertical distribution. Although
Norwegian populations migrate into the subtidal in winter (Lein 1980), it is generally intertidal in Europe. In New England it extends into the subtidal all year round,
presumably due to the absence of predators and competitors (Vermeij 1978).
Species that cooccur with L. littorea in New England and lay benthic egg cases may suffer greatly reduced reproductive success where L. littorea is abundant, but
there is no evidence yet to suggest competitive exclusion of other grazers by L. littorea. Behrens Yamada & Mansour (1987) contrast the competitive assymetry
between L. littorea and L. saxatilis with the well matched Littorina species pairs of the North American west coast. They suggest that this may account in part for the
success of L. littorea in New England, but this ignores the fact that these two species also cooccur in Europe, where L. littorea appears to form a much more
restrained component of the intertidal system (although see Lein 1980).
The longterm effects of the expansion of L. littorea on soft sediment communities are considered above. But it seems probable that New England intertidal
communities were not heavily influenced by epilithic grazers before the advent of L. littorea and it is unclear how much this has changed them since the mid nineteenth
century. Vadas & Elner (1992) speculate on this. Algal blooms following L. littorea harvesting in New England are suggested as giving insight into conditions before
the introduction of L. littorea. This takes no account of the historic component of community structure; removing an animal may result in quite different conditions from
those prevailing before its introduction. Vadas & Elner (1992) suggest a longterm change on sheltered shores as intense grazing by L. littorea may reduce recruitment
success of Ascophyllum nodosum, which is a major space occupier. Lastly they also suggest that use of drift algae by Littorina littorea may have driven the sea
urchin Strongylocentrotus droebachiensis to exert greater grazing pressure on attached algae, thus extending the indirect influence of Littorina littorea into the
sublittoral.
Barker & Chapman (1990) point out that generalizations about grazer/algal interactions on rocky shores that have been derived from the study of L. littorea in New
England are based on an abnormal situation: that is, a community which results from the explosive population expansion of an invasive species rather than a coevolved
community. Underwood et al. (1983) argue against the probability of communities coevolving where most species have dispersive larvae, because of the
unpredictability of species densities, interactions, or even presence or absence. However, there is evidence for a longterm trend of change in the New England rocky
intertidal that is absent from systems not subject to invasive species.
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Detritivores
Littorinids living on angiosperms mostly feed on epiphytes or dead material, rather than live angiosperm material (e.g. Robertson & Mann 1982, Kohlmeyer & Bebout
1986, Bebout 1988, cited in Barlocher et al. 1989a, Barlocher et al. 1989b). On nonwoody plants epiphytes can have a strong inhibiting effect on photosynthesis
through shading (SandJensen 1977), so that epiphytic grazing can be beneficial to the host plant (Williams & Seed 1992, see below).
In the case of littorinids living on woody plants (e.g. mangroves), research has focused mainly on patterns of zonation (see references in Reid 1986) and colour
polymorphism (see McQuaid 1996). There has been very little work on their community effects and little is known about grazing. One recent study emphasizes the
importance of dead plant cells and fungi and mangroves may act as a physical substratum and source of detritus, rather than as a primary food source (Kohlmeyer &
Bebout 1986).
Littoraria irrorata
The saltmarsh snail Littoraria irrorata (referred to throughout the literature as Littorina irrorata) is an extensively studied example of a vegetation dwelling littorinid.
Littoraria irrorata is a highly integrated component of saltmarsh communities and very important in recycling of both energy and nitrogen through the system. Temkin
(1983) estimated that the oldest snails in a 28yearold saltmarsh in Virginia were the same age as the marsh itself.
Climbing behaviour An inescapable aspect of the ecology of L. irrorata is climbing. At least during summer (Vaughn & Fisher 1992), L. irrorata has a circatidal
pattern of vertical movement between leaves and stems above the water level during high tide and the base of plants or the sediment surface at low tide. This behaviour
affects species interactions and has been linked with predator avoidance (see below). However, we must assume that sediment feeding is important. Studies on
climbing behaviour have invariably asked why L. irrorata climbs up plants. We could equally ask why it climbs down.
The orientation of L. irrorata towards stems of the angiosperm Spartina alterniflora seems to be by visual cues. L. irrorata appears to have better vision than
most gastropods (Hamilton & Winter 1982, Hamilton et al. 1983) and orientates towards visual cues both in the laboratory and in the field. When placed on the
sediment on a rising tide it shows nonrandom movement towards the nearest plants and will orientate towards black rods held in its vicinity rather than transparent
ones (Hamilton 1977a). On the open sediment it will also orientate towards dark areas on its horizon. These presumably simulate the presence of vegetation and
animals move either up or downshore towards them (Hamilton 1978).
L. irrorata follows conspecific mucous trails (Hall 1973, Hamilton 1977b), presumably using the cephalic tentacles to do so (Peters 1964). Mucous trails have
polarity (Gilly & Swenson 1978, Raftery 1983) and L. irrorata can infer the direction of movement of the snail that laid down the trail for at least 60min afterwards
(Stirling & Hamilton 1986). This may also be important in orientation towards Spartina leaves. Kemp et al. (1990) found that snails in experimental enclosures with
Spartina were more likely to climb the enclosure walls (rather than the plants) if either the plants had not been used by snails for some days previously or dead leaves
had been removed. Either treatment would remove mucous trails from the plants. An interesting finding by Tankersley (1990) is that snails will follow a trail more readily
if the snail that laid it down was reacting to a visual cue. Thus snails are more
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likely to follow other snails that are orientating towards plants.

Climbing up Spartina could be related to avoidance of physical and/or biological hardship. Climbing avoids submersion, but as Littoraria irrorata can survive over
6 wk under water it is clearly not related to the danger of drowning (Bleil & Gunn 1978). McBride (1986) found that, at high temperatures, respiration was greater in
water than in air. This may be because submerged animals cannot use evaporative cooling and experience higher body temperatures. McBride suggests that climbing
may therefore minimize respiratory stress. McBride et al. (1989) also found that snails sometimes climb at low tide and suggest that this may avoid thermal stress.
McQuaid et al. (1979) showed that body temperature of the climbing terrestrial snail Theba pisana decreased with height above the ground and that survival was
greater on plants than on the ground. Climbing behaviour in both this species and Littoraria irrorata can be triggered by high temperatures in the laboratory and in the
field (McQuaid et al. 1979, McBride et al. 1989). Thermal stress could readily explain Warren’s (1985) finding that snails held close to the substratum, but protected
by cages from predators, showed higher mortality in summer than snails well above ground level.
Predation A more frequently examined reason for climbing is predator avoidance. L. irrorata is generally very abundant in saltmarshes, and although some predators
such as the fish Fundulus heteroclitus do not feed on it (Kneib & Stiven 1978) many other predators do. There are a few reports of bird predation on Littoraria
irrorata (e.g. Wilber & Herrnkind 1984) but the major predators are all aquatic. Among these the blue crab Callinectes sapidus and the conch Melongea corona
are especially important. Hamilton (1976) found that blue crabs swim to the surface of the water at high tide and pluck off snails at or near the surface. Conchs do not
take snails out of water, but eat any that fall in or that are submerged on short plants. Hamilton made the obvious inference that climbing was a form of predator
avoidance. Although Stanhope et al. (1982) disagree, there have been a number of studies showing that climbing reduces predation. Blue crabs select intermediate
sized snails (Hamilton 1976, Stanhope et al. 1982, West & Williams 1986, Vaughn & Fisher 1988) and Stanhope et al. found that climbing too is partially size
dependent. The size classes of snails most vulnerable to crab predation were also more likely to climb, but only about 25% of the population left the water at high tide.
Stanhope et al. also found that the addition of crabs to experimental quadrats did not elicit any increase in the number of snails leaving the water.
Predator exclusion experiments, however, demonstrate a significant reduction in predation for snails that are allowed to climb above the waterline at high tide
(Warren 1985, Vaughn & Fisher 1988). Warren found that more snails were eaten by conchs than crabs. Both conchs and crabs can move faster than Littoraria
irrorata and the climbing response entails avoidance of predators, rather than escape from them (Warren 1985). This could explain the failure of crabs to elicit an
escape response in the field. To allow avoidance, climbing must be cued by factors other than the presence of predators, for example rising tides.
Sizedependent prey selection by crabs may control both the size structure of L. irrorata populations (Stiven & Hunter 1976, Vaughn & Fisher 1988) and seasonal
vertical migration of small snails on the shore (Temkin, 1983). The abundance of intermediate sized snails tends to be negatively correlated with the abundance of crabs
that recruit in the spring (Dudley & Judy 1973) and are more numerous low on the shore (West & Williams 1986). Predation rates by crabs are also highly seasonal
(Warren 1985). Because naïve crabs are inefficient and need to learn to attack snails (Stanhope et al. 1982, Warren 1985), both mortality and injuries caused by
unsuccessful attacks are more frequent in spring (Warren 1985). Although small snails are completely crushed by crabs, larger snails often survive attacks, as
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shown by scars on repaired shells. Stanhope et al. (1982) suggest that 60% of attacks may be unsuccessful. When subject to crushing, shells tend to crack along
growth lines or sutures. Areas of repair do not weaken the shell and individual snails may bear the scars of several attacks (Stanhope et al. 1982, Blundon & Vermeij
1983).
Conchs attack L. irrorata by pushing the operculum aside and it is difficult to infer their success rate, but Warren (1985) found that more snails were killed by
conchs than by crabs in all seasons. Certainly L. irrorata shows a strong escape response to contact with conchs, or even their mucus (Dix & Hamilton 1993). The
fact that conchs do not damage the shell during feeding is important to hermit crabs, particularly Pagurus longicarpus.
Wilber & Herrnkind (1984) found that fresh prey shells from feeding conchs were the primary source of shells for P. longicarpus in a saltmarsh in Florida. There
are many shells available on the sediment surface, but their quality decreases with time and feeding conchs provide a source of high quality shells. McLean (1974)
recorded the attraction of hermit crabs to feeding sites and Rittschof (1980a,b) found that this was due to chemical cues provided by small molecules released by
injured prey. Hermit crabs begin to arrive at these sites within 1–3h (Rittschof 1980a) and establish a dominance hierarchy (Wilber & Herrnkind 1984). The dominant
crab usually has the first opportunity to use the empty shell released by the predator and release of its old shell causes a cascade of very rapid exchanges among the
remaining crabs. As each individual obtains a better shell it leaves the area (Rittschof 1980a). Better shell fit improves predator resistance (Vance 1972), growth and
fecundity (Bach et al. 1976, Fotheringham 1976) of hermit crabs. For example, mangrove hermit crabs take the shells of Littorina varia in greater proportions than
their availability, but do not use the shells of L. fasciata at all. This correlates with the defensive properties of the shells (e.g. strength) and the results of vulnerability
experiments (Borjesson & Szelistowski 1989). So hermit crab populations are likely to benefit not so much from high densities of Littoraria irrorata as high rates of
conch predation. Pagurus longicarpus is thought to transport Littoraria irrorata shells subtidally so that even subtidal hermit crab species benefit from heavy conch
predation (Wilber & Herrnkind 1984).
Feeding Our understanding of the trophic position or role of Littoraria irrorata in saltmarshes has undergone several changes with time. It is found on soft sediments
during low tide and on the upright blades of Spartina at high tide (Hamilton 1976). The early perception seems to have been that feeding occurred on the sediments,
while Spartina shoots were used solely to escape subtidal predators. Teal (1962) divided primary consumers in Spartina marshes into two groups; those that fed
directly on Spartina and those (including Littoraria irrorata) that derived their energy from algae and detritus. Marples (1966) supported this with his finding that L.
irrorata took up radioactive tracer (P32) that had been sprayed on the sediment, but not that which was injected into Spartina stems. Tracer uptake from sediments
can be misleading as Zn65, for example, can be taken up through the foot, without feeding (Ireland 1983). Haines (1976) later found that the stable carbon isotope ratio
of Littoraria irrorata is similar to that of Spartina.
Alexander (1979) finally looked inside the animal and found that dead Spartina formed 90% of gut contents. About 40% of Littoraria irrorata fed on sediments at
low tide and only 4% used live Spartina; there is no difference in isotope ratio between live and dead Spartina. Stiven & Kuenzler’s (1979) finding that altering
Spartina detritus density did not affect growth or mortality of Littoraria irrorata seems anomalous, but they manipulated detritus density by adding cut plants to
experimental enclosures. Under natural conditions dead Spartina leaves remain attached and little material is found on the sediment surface
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(Schubauer & Hopkinson 1984). Littoraria irrorata feeds primarily on standing dead Spartina (Kemp et al. 1990) so that the addition of cut material may not have
directly altered the availability of preferred food.
Alexander (1979) noted that Spartina material was unaltered in the faeces, but diatoms, fungi, bacteria and protozoans on the Spartina and sediments were
assimilated. Preconditioning of angiosperm detritus by microorganisms and fungi is required by many deposit feeders (Lopez & Levinton 1987) and fungi are important
in the metabolic transformation of standing dead Spartina. Littoraria irrorata feeds on the mesophyll of dead leaves, rather than on their surfaces, and so takes in
large quantities of fungi (Kemp et al. 1990). Colonization of dead Spartina by fungi increases the availability of proteins to Littoraria irrorata (Barlocher et al. 1989a)
as it is capable of digesting fungi (Bebout 1986, cited by Kohlmeyer & Bebout 1986). L. irrorata is also capable of breaking down the primary components of
angiosperm cell walls, such as cellulose and xylan (Barlocher et al. 1989b), but does not grow on a diet of recently dead leaves that have not been colonized by fungi
(Barlocher & Newell 1994).
Kemp et al. (1990) suggest that L. irrorata may have an important role in detritus cycling by converting standing dead Spartina into fine particulate matter on the
sediment surface. They give a preliminary estimate that Littoraria irrorata may ingest roughly 13–27% of Spartina production per year, but consumption is highly
seasonal. Growth (Temkin 1983) and respiration (Shirley et al. 1978) both decrease in winter and Kemp et al. (1990) indicate that ingestion of dead Spartina could
be 100% in summer and early autumn. This could be exacerbated by the fact that sediments are not used for feeding in June (Alexander 1979).
In summary, Littoraria irrorata has not been shown to have direct effects on the distribution of other species (although there has been no work on this) but it is an
important prey species for conchs and crabs and may have indirect effects on hermit crab populations. More important is the role of L. irrorata in nutrient cycling
within saltmarshes, making Spartina detritus available to other species.
Epiphytic grazers
Relatively little work has been done on these species, but grazing of epiphytes on nonwoody angiosperms or macroalgae can have significant effects because epiphytic
shading can reduce macrophyte photosynthesis, for example, Littorina in Nova Scotian Zostera marina meadows. I shall use the name L. neglecta from the original
paper, though this may be wrong (see McQuaid 1996). These populations can reach enormous densities (45000m−2) and influence photosynthetic rates of Zostera by
controlling the amount of epiphytes and detritus on Zostera leaves (Robertson & Mann 1982). Because angiosperms derive nutrients from the sediments, epiphytic
grazers such as Littorina neglecta may also play an important role in nutrient cycling. Grazing recycles nutrients from the periphyton via faeces, but L. neglecta also
feeds on bluegreen algae, which occur on some plants and on the surface of the mud. These fix nitrogen rapidly and again grazing makes this available to Zostera
(Robertson & Mann 1982). There is no suggestion that this is important for macroalgae, which obtain nutrients from the water column.
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Littorina mariae
The best studied example of an epiphytic grazer is probably Littorina mariae. Unfortunately L. mariae was for many years confounded with the macroalgal grazer L.
obtusata as a single species, L. littoralis. It is convenient initially to consider the two species together.
Studies on possible links between colour morph frequency and predation of L. obtusata and L. mariae by fish are summarized in McQuaid (1996), but there is a
further suggestion that predation by crabs may influence niche separation between these two species. Reimchen (1982) examined L. obtusata and L. mariae from a
large number of sites in the British Isles and correlated size with wave exposure. The two species diverge in size under more sheltered conditions where L. mariae tend
to be smaller than conspecifics on exposed shores and L. obtusata larger. Crab densities are also higher on sheltered shores and Williams (1990a, 1992b) suggests
that niche separation between the two may be largely driven by predation. He suggests that the lower limits of L. obtusata are set by crab predation and that L.
obtusata has a refuge in space from crab predation by occurring on the mid shore. Williams also suggests that L. mariae may have a refuge in time, having an
essentially annual life cycle with adults being most abundant in winter when crabs move offshore and predation is least intense.
The zonation of the two species reflects their preferences for different algae. The vertical distribution of L. obtusata encompasses that of L. mariae, which is more
restricted. L. mariae occurs mainly (sometimes entirely) on Fucus serratus and is virtually confined to the low part of the shore where F. serratus occurs. Janke
(1990) found that the population of Littorina mariae fluctuated with that of Fucus serratus, but that protecting Fucus fronds from wave action allowed an increase in
Littorina mariae density, suggesting that water movement and rubbing of fronds may normally limit densities. L. obtusata occurs over a much wider area of the shore,
but is most abundant on the midshore canopy of Fucus vesiculosus and Ascophyllum nodosum (Watson & Norton 1987, Williams 1995). Underwood (1972),
possibly working with a combination of the two species as Littorina littoralis, found that, when placed on the bottom of a tidal tank, the snails climbed to the top
unless they encountered fucoid algae (Fucus spp. or Ascophyllum nodosum) part way up, in which case they stopped when they reached the algae. Williams (1995)
suggests that movement of transplanted animals of both species towards their original zones may be mediated by attraction to their respective host algae.
In Scotland, the adults of both species are primarily confined to the fronds of canopy algae in the Fucus zone while juveniles also occur on rocks and ephemeral
algae such as Ulva (Watson & Norton 1987). Both species have radulae that are blunter than those of rockdwelling littorinids (Watson & Norton 1987, Steneck &
Watling 1982) and they differ from each other mainly in the cusps of the teeth, the radula of Littorina obtusata apparently being better designed for excavating
macroalgae.
L. mariae faeces indicate that it feeds mainly on epiphytic diatoms and detritus on the surface of Fucus serratus. We have seen above that epiphytic grazers not
only change the biomass of diatoms, they also change the species composition. For example, F. serratus grazed by Littorina mariae carries a periphyton of larger
diatom species than ungrazed plants (Williams 1992a). Williams (1990b) argues that as algal surfaces are renewable, the constraints they impose on grazers are
different from those imposed by rock. This is reflected in the link between life cycles of grazer and host, for example between Lacuna pallidula and Fucus serratus
and between Littorina mariae and Fucus serratus (Williams & Seed 1992). Also interactions between epiphytic grazers and algae are different from interactions
between algal grazers and algae. Epiphytes are often, although not always, detrimental to
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algae and so grazers of epiphytes can be beneficial to the host alga. The epiphyte load on algae is reduced where grazers are present and the distribution of epiphytes
on different parts of a plant is negatively correlated with that of grazers (Williams & Seed 1992). Some authors have suggested that, by removing epiphytes, grazers can
actually improve the fitness of host plants (Lubchenco 1983, Williams 1992a). D’Antonio (1985) found that epiphytes reduce growth, reproductive output and increase
the probability of breakage of the red alga Rhodomela larix. Amphipods reduce these negative effects by decreasing the cover of epiphytes and D’Antonio suggests
that Littorina scutulata living on Rhodomela larix and feeding mainly on epiphytic diatoms may also have a similar effect by reducing diatom cover. Williams (1992a)
suggests that epiphytic grazing may affect lowshore community structure by enhancing the success of host algae.
Macroalgal grazers
Algal defences against grazing

Clearly algal defences evolved in response to grazing by a range of herbivores, in some cases including littorinids, but in this section I emphasize the link between
littorinids and such defences. Where this link is close, it may reflect the response of littorinids to particular defences as much as the evolution of defences in response to
littorinid grazing.
Macroalgal defences against grazing may influence the outcome of species interactions and take three main forms. First, many species that are highly palatable (e.g.
microalgae and ephemeral macroalgae) both grow and propagate very rapidly. The unpredictable appearance of ephemerals in time or space prevents macroalgal
grazers from specializing on them. They are often eaten by generalist herbivores, and adult plants represent sporelings that have escaped grazing. This emphasizes
survival of the species rather than of individual plants. Individual plants generally have no defence against grazers, and are shortlived with continuous, high availability of
propagules. Later successional, perennial species tend to sacrifice rapid growth for grazer resistance (Littler & Littler 1980) and have structural or chemical defences
that are often negatively correlated with the grazing preferences of grazers, including littorinids.
Obvious structural defences include calcification, branching patterns and a crustose growth form. Particular growth forms such as coarsely branched coralline species
are comparatively resistant to grazers (Padilla 1984) although calcification does not increase the force required by limpets to remove tissue (Padilla 1989). While some
measure of resistance to puncture or abrasion is often negatively correlated with feeding preferences among upright algae, grazers such as littorinids do not always eat
those crusts that are mechanically easiest to remove (Dethier 1994).
A crustose growth form can be a particularly effective defence against grazers. Bertness et al. (1983) examined not just physical properties of algae, but also their
effects on grazers. While a mechanical measure of scratch resistance was greatest in the foliose Chondrus crispus because of its thick cuticle, tooth wear in Littorina
littorea was severe only when it fed on crustose algae. Bertness et al. (1983) linked this to the arrangement of cells in the thallus. They suggest that many small cells
packed closely together offer greater resistance to radular damage than a thick cuticle covering fewer, large cells. Watson & Norton (1985b) also use cell size (which
influences the cell wall:cytoplasm ratio) to explain differences in toughness in different parts of the same foliose plant. Slocum (1980) showed that the
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crustose phase of Gigartina papillata increased in abundance in the presence of limpet and littorinid grazing, while the blade phase decreased. Similarly Lubchenco &
Cubit (1980) correlated the occurrence of different phases in heteromorphic algae to the seasonality of grazing pressure. They showed that exclusion of Littorina
littorea resulted in algae appearing as upright forms in seasons when they would normally occur only as crusts.
While it is convenient to refer to “crustose species” as a functional group of algae, they actually show a wide variety of strategies for resisting grazers, including
chemical defence (Dethier 1994). Likewise a crustose form may work against some epilithic grazers such as L. littorea and L. scutulata, but not against others such as
L. sitkana, perhaps because of differences in tooth structure (Behrens Yamada 1992, Dethier 1994).
The effect of chemical defences on feeding preferences can be demonstrated using homogenates solidified with agar as this removes differences in texture, etc. This
method was used by Geiselman & McConnell (1981). They found that the results for homogenates correlated with feeding preferences in the field. Chemical defence of
noneaten algae was achieved, as in terrestrial plants, by a mixture of polyphenols.
There are a number of other correlations between grazing preferences and polyphenol levels in algae. Van Alstyne (1988, 1990) showed that survival of Fucus
distichus was not correlated with the density of littorine grazers (Littorina scutulata and especially L. sitkana). However, injured plants produced more polyphenols
and were less heavily grazed under natural conditions. Similarly, littorines preferred meristematic tissue to adventitious branches that had a higher polyphenol content
(Van Alstyne 1989). Among Fucus spp., F. distichus suffers lower rates of grazing than others and has particularly high levels of polyphenols (Barker & Chapman
1990). These authors hypothesize that Fucus species in areas with high grazing pressure channel more resources into antiherbivore defences. This makes sense
because a high phenolic content can be induced, but Yates & Peckol (1993) impose a very important qualification. Like Van Alstyne, they induced a rise in polyphenol
content in some populations of F. vesiculosus by simulating grazing damage and found that Littorina littorea preferred phenolicpoor to phenolicrich Fucus
vesiculosus. However, polyphenol content was consistently higher at a lownitrogen site than at a highnitrogen site and a rise in phenols could only be induced in F.
vesiculosus growing under lownitrogen conditions. In other words, polyphenol content is affected by both grazing level and abiotic environmental factors and there is
interaction among these effects.
Orians & Janzen (1974) observe that any mechanism for reducing predation of eggs is unlikely to be effective against all predators. The same argument applies to
grazers. Structural defences are effective against generalist macroalgal grazers. Polyphenols also deter generalist grazers, but some grazers have the specialized ability to
cope with them. Chemical defences seem to allow the coevolution of specialist grazers that are immune to the defences of a particular alga. Littorina obtusata is
attracted to and eats Ascophyllum nodosum (Watson & Norton 1987) despite the high phenol content shown by Geiselman & McConnell (1981). We shall see later
that, among specialized grazers, different populations of the same species may eat different algae.
Littorina obtusata and the genus Lacuna offer good examples of intertidal and, for some Lacuna spp., subtidal macroalgal grazers. The biology and community
effects of macroalgae grazers seem to be quite different to those of epilithic species. This has been shown particularly clearly in recent work in Canada on Littorina
obtusata, which contrasts it with L. littorea. The primary difference is that the epilithic grazers which have been studied influence the distribution and density of adult
algae by grazing sporelings while those feeding on macroalgae influence the canopy and survival of mature plants.
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Littorina obtusata
The importance of predation to the population structure of L. obtusata varies geographically. The results of experiments on crab preferences for different size classes
of L. obtusata differ for areas where L. mariae does and does not occur. Rangeley & Thomas (1987) worked with Carcinus maenas from Canada, where Littorina
mariae does not occur. In the laboratory Carcinus was less successful in attacking small Littorina obtusata (although size classes were not defined) because of
difficulty in manipulating them. The mean size of snails that survived was smaller than of those that were eaten. Difficulty in handling small snails was presumably not due
to naïvety of the crabs as they were collected from shores where L. obtusata is found. In Britain, where the two snail species cooccur, Reimchen (1982) found the
opposite: large L. obtusata were more difficult for Carcinus to open; Littorina obtusata, from shores where crab densities are high, have more shell injuries and tend
to be larger (see also Williams 1994). Reimchen found that even crabs substantially larger than those used by Rangeley & Thomas could manipulate small snails (6mm
shell width) and in choice experiments preferentially ate the smallest prey. Reimchen makes no mention of crabs having difficulty handling small snails. Certainly shell
thickness is important in determining how vulnerable snails are. Seeley (1986) found that L. obtusata in New England had shorter spires and thicker shells subsequent
to the introduction of Carcinus maenas to New England and in areas where crab densities were high. She also demonstrated greater mortality among tallspired, thin
shelled individuals when snails were tethered in areas of high crab density.
Littorina obtusata feeds mainly on adult fucoids, preferring these to Ulva or Fucus sporelings, (Hawkins & Hartnoll 1983, Watson & Norton 1987, Williams
1990a) and is attracted to fucoids rather than understorey species (Watson & Norton 1987). Littorina obtusata is most strongly attracted to Fucus vesiculosus and
feeds preferentially on it rather than other fucoids (Watson & Norton 1987, Barker & Chapman 1990), making small holes through the fronds (Viejo & Arrontes
1992, who use the name Littorina littoralis). This is reflected in the hydrogen isotope ratio of the snail, which is the same as that of Fucus vesiculosus, but not of
coexisting algae, such as Ulva and Chondrus crispus (Estep & Dabrowski 1980). Littorina obtusata also occurs on and eats Ascophyllum nodosum, despite its
high polyphenol content, although it has a low preference in food choice experiments (Watson & Norton 1987).
Watson & Norton (1987) suggest that the main ecological impact of grazing by littorinids lies in their effect on sporeling survival and, in some cases, in their
preference for reproductive tissues. They suggest that grazing effects on adult macroalgae may be the least significant of their activities and cite the example of
Lubchenco’s work on Littorina littorea. However, L. littorea is an epilithic grazer and manipulative field experiments by Chapman (1989) have clearly brought out a
direct contrast with the community effects of macroalgal grazers. Perhaps the most important point is that, unlike L. littorea, L. obtusata does not facilitate succession
as it prefers adult fucoids to sporelings or ephemeral algae (Chapman 1989, 1995).
Lubchenco (1983) suggests that periwinkles (including L. obtusata) remove epiphytes from Fucus vesiculosus and so may be beneficial to the plant by decreasing
drag. But Littorina littorea, like Nodilittorina unifasciata, mainly affects adult algae indirectly by grazing sporelings and so influencing recruitment. In Britain,
Littorina obtusata feeding on Ascophyllum nodosum physically damages the plant, but does not appear to affect the success of individual plants or the overall
population of the alga. The result is quite different when it is maintained on Fucus serratus. Littorina obtusata removes the fronds of Fucus serratus entirely and
reduces the plant to the stipe and midrib (Williams & Seed 1992).
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In Nova Scotia, Chapman and his coworkers have been studying communities that include four species of fucoid algae. These are Fucus distichus in highshore
pools and, in descending order down the shore, F. spiralis, F. vesiculosus, F. evanescens. They have found that grazing effects vary according to three criteria: the
species of fucoid, the zone on the shore, and whether one considers mature plants or sporelings. Considering sporelings, Chapman & Johnson (1990) found that
grazing had little effect on recruitment low on the shore, and suggested that grazing effects for each species of alga increased upshore. Interzone differences in grazing
are likely to be due to the confounding effects of differences in both algal growth rates and which grazers are present. Apart from amphipods, these are mainly
littorinids: Lacuna vincta, presumably on the low shore, Littorina obtusata on the mid and low shore, L. saxatilis higher upshore and L. littorea in highshore pools
and low on the shore (Chapman 1989, Chapman & Johnson 1990). Littorinids are also among the most abundant animals in pools, with different species predominating
in different zones (Metaxas et al. 1994). Thus different types of grazers occur at different heights on the shore.
In highshore pools, littorinid grazers (mostly L. littorea) prevented the establishment of macroalgae (Parker et al. 1993). While grazers (including L. littorea and
amphipods) strongly influenced survival of Fucus distichus sporelings, they did not affect the canopy of mature plants. Herbivory was more important to sporelings
than canopy shading and the presence or absence of a canopy only significantly affected their survival in the absence of grazers (Chapman 1990). In the next zone
down (the upper eulittoral), grazers (mainly Littorina obtusata and L. saxatilis) reduced the canopy of Fucus spiralis, but not sporeling survival. However, by
reducing the canopy they may influence sporeling survival indirectly (Chapman 1989).
In rather different studies of primary succession following massive ice scour of the lower shore where F. vesiculosus and F. evanescens occur, McCook &
Chapman (1991, 1993) found that littorinids were restricted to crevices so that much of the substratum was out of their reach. In this case herbivory did not mediate
plantplant competitive interactions and was not an important factor in succession. Instead succession was driven primarily by patterns of spore settlement. On the other
hand, the density of Littorina obtusata did increase with greater canopy cover as it congregated under the algae, as do limpets (Hartnoll & Hawkins 1985).
We have already seen that, under many circumstances, littorinids can mediate competitive interactions between fucoid and ephemeral algae. In several studies they
have been found to facilitate fucoid growth by removing ephemerals (Lubchenco 1978, 1983). However, littorinid grazing can also enhance the success of ephemerals
(Chapman 1990, Chapman & Johnson 1990). One mechanism for this is suggested by Santelices et al. (1983). They found that ephemeral algae regenerated more
frequently from grazer (sea urchin) faeces than did late successional algae. Regeneration from the faeces of L. peruviana was lower than for other mollusc grazers
(Santelices & Correa 1985), but this could reflect the background of the animals used, as the efficiency of digestion in littorinids is influenced by feeding history (Abele
Oeschger & Theede 1991). Breeman & Hoeksema (1987) suggest that resistance of digestion and regeneration from L. littorea faeces can be an important
mechanism for propagation of the red alga Rhodochorton purpureum.
The effects of littorinid grazing on community structure are further complicated because, not only can ephemeral algae compete with fucoids (Lubchenco 1986), but
they may also facilitate their growth (McCook & Chapman 1993). McCook & Chapman conclude that where herbivory is patchy the effects of inhibition or facilitation
of Fucus growth may also be patchy.
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Perhaps the most important point that emerges from the work of McCook & Chapman (1993) and that of Barker & Chapman (1990) is the need to refrain from
generalizations based on interactions between pairs of species. It is not reasonable to assume that Littorina affects Fucus in particular ways. Different species of
Littorina interact differently with different species of Fucus. This is brought out very clearly in the following section on Lacuna. Underwood and his coworkers have
gone further in showing that particular species interactions also vary from place to place and from time to time in the same area (Underwood et al. 1983).
Lacuna spp.
Little has been published on the effects of predation on Lacuna, beyond the comment by Johnson & Mann (1986) that the populations they studied were not limited by
predation.
The genus Lacuna is a good example of different species eating different algae and producing quite different consequences. Some Lacuna species have been shown
to coexist with food algae without causing mortality and causing only minor damage or loss of biomass, while others have been shown to have devastating effects on
food algae. When these are canopy species the structure of the entire algal community may be expected to be affected. The effects are strongly influenced by details of
the biology of both grazer and alga as well as the specific interaction between the two. Obviously the part of the plant that is grazed is important and may reflect the
morphology and chemical composition of the alga, but feeding sites are also a function of species pairs. L. vincta and L. parva feeding on Phycodrys rubens attack
different parts of the plant (Ockelmann & Nielsen 1981). Likewise, the type of damage done is important and depends on the food species as well as the behaviour
and radula structure of the grazer. But other aspects are equally important, such as population dynamics of the snail (Lacuna often shows abrupt population explosions)
and the seasonality of grazing (which affects recovery of the plant or the response of understorey species to removal of canopy).
Even within one species behaviour may differ dramatically. According to Ockelmann & Nielsen (1981), British and French populations of L. parva live on
Chondrus crispus, which Danish snails will not eat, even when starving. Not surprisingly, completely different effects have been shown for the same species of Lacuna
in different studies.
Lacuna species have annual life cycles and lay egg masses that are attached to macroalgae and may, like L. vincta, have planktonic development or, like L.
pallidula and L. parva, have direct development, with crawling juveniles hatching from the benthic eggs (Smith 1973, Fretter & Manly 1977, Grahame 1977, 1982,
Ockelmann & Nielsen 1981, Southgate 1982). Annual life cycles may be closely linked to the life history of the host alga (Smith 1973, Grahame 1985) and result in
marked seasonality of recruitment and of grazing pressure (e.g. Trowbridge 1992).
Veligers of L. vincta will settle and spawn on a number of macroalgae, including red algal turfs (Fretter & Manly 1977, Thomas & Page 1983) and juveniles (under
1mm) feed on diatoms and detritus, possibly because the radula cannot penetrate macroalgae (Fretter & Manly 1977, Smith et al. 1985). Adult L. vincta are generalist
macroalgal grazers and will feed on a variety of brown algae. Some authors have found that grazing by adult L. vincta can damage individual plants quite severely but
either make no mention of obvious repercussions on the rest of the community (Norton 1971, Smith 1973), or specifically comment on the absence of such effects
(Grahame 1985, Johnson & Mann 1986). Others record the devastating effects of overgrazing (Fralick et al. 1974, Thomas & Page 1983). There is no
Page 290
biogeographic trend in these results and each of the studies considered different species of algae. Perhaps the most significant fact is that the effects of grazing do not
appear to depend solely on the density of snails, but rather reflect interactions between particular species pairs. It is difficult to compare densities in these various
studies as they are measured in different ways (density m−2, density plant−1, density gram of plant−1, etc.), and the values given often refer to different size classes that
may be eating different things. Still, the independence of effects from snail density can be seen in a comparison of the findings of Johnson & Mann (1986) with those of
Thomas & Page (1983).
The latter authors observed densities of up to 1570m−2 L. vincta feeding on low intertidal Fucus edentatus in the Bay of Fundy. These animals were 3–5mm long
(possibly they migrated upshore from the subtidal) and caused weight losses of up to 79% in F. edentatus. Grazing also caused a change in growth form. Grazed
plants became thicker, rather than elongating, but were frequently reduced to midrib tissue. They noted that species which are grazed by Lacuna vincta were normally
low in abundance on the shore. Similarly, Fralick et al. (1974) found that subtidal plants of Laminaria saccharina and L. digitata in New Hampshire had badly
damaged blades and were reduced to holdfasts and stipes in areas of particularly heavy Lacuna vincta grazing, even though the animals were less than 2mm in length.
Johnson & Mann (1986) give four major reasons for L. vincta being a potentially important grazer: it is highly dispersive even after larval metamorphosis (see Martel
& Chia 1991a,b); its numbers are not limited by predation in their study area; and it can attain very high densities; it feeds preferentially on the dominant canopy
species. They found that after a pulse of recruitment, small L. vincta (0–2mm long) reached densities of 4500m−2 in a Nova Scotian kelp bed, and even when 0–4mm
long, mean densities were around 2000m−2. Despite such high densities L. vincta grazing reduced only the canopy of Laminaria longicruris, and not its survival.
Parts of the plants were avoided either because they had a high phenol content, were particularly tough, or had low nutritional value and grazing was concentrated on
the frilly margins of the blades. Grazing removed only about 0.05% of L. longicruris biomass, although it did promote tissue loss through wave action, by weakening
the blade edges. Although Lacuna reduced canopy shading by Laminaria, under storey algae were unable to take advantage of this because of the seasonality of
grazing intensity. Grazing decreased towards autumn due to enormous mortality of Lacuna, and the canopy recovered before most understorey species entered their
spring growth phase.
Obviously the availability of algae is important and may affect grazing behaviour. Ockelmann & Nielsen (1981) found that L. parva in the laboratory fed on the
thinnest, youngest parts of Phyllophora pseudoceranoides and Delesseria sanguinea. Even when these became scarce they ate the ribs and older parts but avoided
other algae. Similarly, Johnson & Mann (1986) found that Lacuna vincta colonized algae other than Laminaria longicruris only when their densities were very high.
Research on other species of Lacuna has noted coexistence with food algae, without comment on dramatic population or community effects through overgrazing,
although individual plants may be badly damaged. This includes L. pallidula feeding on Fucus serratus (Smith 1973), Lacuna variegata feeding on Sargassum
muticum (Norton & Benson 1983) and Lacuna marmorata feeding on Iridaea cordata (Gaines 1985).
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Summary
The consequences of grazer/algal interactions depend on a variety of factors. Chief among these are the type of grazer and grazing pressure exerted and factors
affecting algal growth rates. The latter include season, height on the shore, habitat (pools or open rocks), and degree of exposure. Many of these factors also influence
algal competitive interactions that are equally important. This is brought out very clearly in studies that compare Littorina littorea grazing effects in pools and on open
rocks; in pools at different heights on the shore; and even between pools or open rocks at the same height on exposed and sheltered shores. We also find that workers
get different results when doing the same experiments at different places or different times (examples are given in Underwood et al. 1983, Vadas & Elner 1992 and
Chapman 1995). Usually this is explicable in terms of differences (sometimes subtle), in physical conditions, the species present, species densities or timing of the
experiment (seasonality or weather effects). A very real problem is that, as Chapman (1995) succinctly puts it, “The more we know, the more tenuous the general
models become. There is a danger that everything revealed will become site specific”.
We know comparatively little about the influence of littorinids on microalgae, but they have little effect on the macroalgal community either at the top of the shore
(where macroalgae are probably excluded by physical factors) or in cases where there is a close association between a specialized grazer and a host plant (as between
L. obtusata and Ascophyllum nodosum). Likewise littorinids have little effect where (like Littoraria irrorata) they eat dead material. Littorinids have their most
dramatic effects on the community when they are either generalist macroalgal grazers (especially if prone to population explosions as is Lacuna), or generalist epilithic
grazers that remove sporelings (e.g. Littorina littorea). In the case of macroalgal grazers, community effects are not strictly density dependent, but are influenced by
the details of interactions between grazer and alga. Also important are grazer preferences with respect to algal competitive hierarchies. Quite different effects emerge if
grazers prefer competitively dominant or nondominant algae.
In the case of epilithic grazers, grazing effects can be very patchy and the importance of grazing is a consequence of the effect of grazers on the recruitment of algae.
This becomes complex as algal recruitment is also affected by interactions among algae, particularly the effects of canopy shading on sporeling survival and competition
between perennial and ephemeral algae. The effects of epilithic grazers depend very much on height on the shore, but they are also strongly density dependent and are
greatly influenced by wave exposure as this affects both algal growth rates (and/or competitive hierarchies), snail densities and grazing efficiency. Algae swamp grazers
where algal recruitment or growth rates are high (e.g. low on the shore or with more exposure), while grazers can control algal abundance, to the extent of excluding
macroalgae, where grazer densities are high. Grazing of dominant algae increases algal species richness while removing subdominants reduces richness. The effects of
grazers are also more pronounced in seasons when algal growth is reduced.
The consensus is that predation may affect local densities of littorinids, but that predation does not limit population sizes. Many field studies record either that rates of
predation are low, or that, while large numbers of littorinids are eaten, they form a small proportion of the total population (e.g. McQuaid 1985, Johnson & Mann
1986, Alerstam et al. 1992, Huxham et al. 1993).
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Conclusions
Ecological interactions are not deterministic and we must recognize that the minutiae of biological interactions can profoundly affect their outcome. Nevertheless, we
appear to be reaching an understanding of the general factors that influence some of these interactions.
1. The factors governing community structure can be usefully divided into abiotic and biotic. Abiotic conditions set the biogeographic and physical framework within
which biotic interactions occur (McQuaid & Branch 1984). When severe enough, abiotic factors assume overriding importance. For example, decreases in algal
biomass in summer in Australia and Hong Kong (Underwood 1981, Williams 1993); the removal of species by ice scour in North America (Wethey 1985,
McCook & Chapman 1991, 1993); the overriding of grazing effects by wave action in Germany (Janke 1990).
2. Physical conditions have some predictability. Information on height on the shore, season, in or out of pools, etc. gives us relative values of physical conditions. It is
hotter in summer, even if we do not know how hot.
3. Algal growth rates are strongly influenced by physical conditions and grazer/algal interactions may have predictability depending on the balance between growth
rates and grazing pressure. For example, grazers can control algal biomass or be swamped by algal growth, depending on this balance. Grazing pressure is affected
by a number of factors, including several that affect algal growth, but for epilithic grazers at least, the most important factor seems to be grazer density. The results of
interactions between epilithic grazers and sessile animals can also depend on the relative densities of the species. Grazers can influence barnacle numbers or vice
versa, depending on initial densities.
4. Densities are strongly influenced by recruitment success and we require an understanding of the factors controlling recruitment (e.g. Underwood & Fairweather
1989). Assuming larvae are available (i.e. a viable adult population occurs within effective larval dispersal range) these occur at two levels: a) factors controlling
whether larvae reach a shore (largely hydrodynamic factors and larval behaviour), and b) factors affecting whether settlers survive to recruit into the adult population
(abiotic and biological factors).
Subtle effects exist at lower levels (microhabitat variability in slope, aspect, etc.) and we see these when we use a reductionist approach, for example to small scale
snail distribution (Chapman 1994). At a broader level, variability in density is due at least in part to recruitment success. What we need is an understanding of density
effects and the control of recruitment.
Acknowledgements
This work was generously funded by an Ernest Oppenheimer Memorial Trust Fund Travelling Fellowship, the Joint Research Committee of Rhodes University and the
Foundation for Research Development. I am particularly grateful to those who read and commented on earlier drafts of the paper: A.R.O.Chapman, P.A.Hulley,
D.Raffaelli and especially G.A.Williams. I am infinitely grateful to Lesley McQuaid for dealing with the references.
Page 293
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PEDUNCULATE CIRRIPEDES OF THE GENUS POLLICIPES

MARGARET BARNES
The Scottish Association for Marine Science, Dunstaffnage Marine Laboratory, P.O. Box 3, Oban, Argyll PA34 4AD, Scotland

Abstract Pedunculate cirripedes, particularly those found on floating objects washed up on the shore, have long been called goose (or gooseneck)
barnacles because of their graceful appearance and the long stalk that resembles the neck of a goose. These barnacles are rarely found growing on
fixed intertidal substrata. The only truly intertidal pedunculate cirripedes belong to the genus Pollicipes of which there are three species. Two are
found on the west coast of the Americas and the third is European. Very little is known about P. elegans from South America. P. polymerrus is
found on the Pacific coast of North America extending from 27°N to 64°N. P. pollicipes is less eurythermal than P. polymerus and has rarely been
found north of the Atlantic coast of France; the most southerly record is about 15°N.
Most of this review concerns P. polymerus and P. pollicipes because of the lack of published work on P. elegans. A general description of the
animals, the arrangement of the calcified plates, the muscles and the nervous system have all been considered. The animals favour exposed habitats
where there is a backwash from surging waves. The strength of the peduncle and its ability to bring the capitulum and the cirri into the best position is
related to the captorial method of feeding. A high haemolymph pressure and the musculature of the peduncle enable it to contract or lengthen
according to the environmental conditions. The peduncle plays a vital part in combating the effects of temperature change and desiccation. It provides
an additional surface for gaseous exchange, and a substratum for settling cyprids.
The size of adults at maturity and their fecundity, spermatogenesis and oogenesis have all been discussed. Fertilization, development of embryos,
and sizes of nauplii have been compared for the species. The ultrastructure of the membranes involved in fertilization and development of embryos has
been examined.
Chemical analyses of bodies, egg lamellae, and calcified structures have been determined. Predation and the effect of pollution have been found to
be important. The commercial exploitation of all three species and its future possibilities have been examined.
Introduction
Pedunculate barnacles and their origin have intrigued naturalists since early times. Ideas that the shells hung on trees by a neck and when shed into the water produced
birds, with beaks like those of geese, originate in myths about their ancestry (HeronAllen 1928, Gurney 1947). The term goose (or gooseneck) barnacle, as a
common name of some stalked barnacles must have originated in this way—as probably did the name barnacle goose.
Barnacles are cirripedes belonging to the Crustacea. They have many forms and a sessile habit. They may be freeliving filter feeders or specialized parasites and can
be found in estuarine or fully marine habitats ranging from the littoral to sublittoral zones extending down to the deepsea vents. Some species are hermaphrodites, with
or without complemental males, while in others the sexes are separate, dwarf males being present in the vicinity of the females.
Attempts at classification of the Cirripedia began even before Darwin but he was responsible for establishing it on a basis that has lasted until the present day
(Darwin 1851, 1854).
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Over the years suggested changes have been made but, in general, it is the system devised by Darwin that has been enlarged and extended as more species have been
discovered and our knowledge has increased (Pilsbry 1907, 1916, Newman & Ross 1976, 1977, Foster 1978, Zevina 1978, Newman 1987, Anderson 1994).
Within the Subclass Cirripedia, the Superorder Thoracica contains the Orders Pedunculata and Sessilia. In the Pedunculata a peduncle is present between the
capitulum and the basal disc that is attached to the substratum (Walker 1992). This Order encompasses, among others, the Superfamily Scalpelloidea, which contains
the Family Scalpellidae with the subfamily Pollicipedinae, and within this subfamily is the genus Pollicipes (Anderson 1994). It is this genus with which the present
review is principally concerned. Darwin (1851, pp. 293–4) in a footnote explains why he decided to use the generic name Pollicipes in preference to Mitella. Pilsbry
(1907) regretted Darwin’s decision and reverted to the earlier generic name of Mitella but, although used by some cirripedologists for several years, it was later once
again replaced by Pollicipes (Newman 1993). This has now become generally accepted. The animals have a peduncle that supports the capitulum. The peduncle is
flexible, provided with muscles and covered with scales. It contains the ovarian tissue. The capitulum is flattened and covered with calcified plates. The capitulum
protects the soft parts of the body.
Darwin (1851) describes six species of the genus Pollicipes but admits that, from the valves on the capitulum, it would be difficult to separate P. mitella from P.
sertus and P. spinosus. The other three species, P. cornucopia, P. polymerus and P. elegans “form one thoroughly natural genus” according to Darwin (1851). In
these last three species the capitulum is covered by regularly formed whorls of plates. The animals also have filamentary processes on the prosoma; these are not
present in P. sertus and P. spinosus (Darwin 1851, Foster 1978). The structure of the maxillule is also different (Foster 1978). Foster (1978) found a complemental
male in P. spinosus; none have yet been reported in other Pollicipes species. He, therefore, suggested that P. spinosus and its synonyms including P. sertus should be
assigned to the genus Calantica as C. spinosa. Pollicipes mitella with its highly sculptured plates on the capitulum and multiarticulated caudal appendages should
remain as a separate genus—probably Capitulum. This leaves Pollicipes pollicipes (P. cornucopia), P. elegans, and P. polymerus as the three species of the genus
Pollicipes (Foster 1978). The European representative has, over the years, been called P. cornucopia or P. pollicipes. The earlier specific name was pollicipes
(Pilsbry 1907) and, in order to avoid confusion, pollicipes will be used in this review (Foster 1978).
The taxonomic authorities for the three species are P. elegans Lesson, P. pollicipes Gmelin, and P. polymerus G.B.Sowerby (Darwin 1851, NilssonCantell
1921). It is unfortunate that a printer’s error (J.B. instead of G.B.) went unnoticed in Barnes & Gonor (1958b) and was perpetuated in a few following papers (Barnes
& Barnes 1959b, Barnes & Reese 1959, 1960).
In many cirripedes the soft parts are protected by a shell composed of various calcified plates and after the animal’s death these plates may be found in marine
sediments. Fossil records of cirripedes have been well documentated by such workers as Hoek (1883), Gruvel (1899, 1902), Karakasch (1903), Joleaud (1910a,
1913, 1916), Broch (1922), Withers (1928, 1935, 1953), Foster (1978), Zevina (1978), Newman (1979a, 1987), Buckeridge (1983), and Foster & Buckeridge
(1987) as well as many others. Broch (1922) described and illustrated the postlarval ontogenetic stages of P. polymerus and Newman (1979a) showed how these
stages could be followed by reference to fossil records. Relics of the genus Pollicipes have been found in Upper Cretaceous sediments (Darwin 1954, Buckeridge
1983,
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Foster & Buckerage 1987, Newman 1993). Foster & Buckeridge (1987) are of the opinion that the separation of the three species of Pollicipes to be dealt with in
this review was caused by the Cenozoic movements of the continents. P. polymerus became isolated in the northeastern Pacific, P. elegans in the tropical eastern
Pacific, and P. pollicipes in western Europe and on the north African coasts of the eastern Atlantic.
When describing Pollicipes several measurements have been used by different workers. These measurements are used on several occassions throughout this review.
They may be defined as follows: CH—height of the capitulum—is the vertical height from the base of the sublateral plates to the tips of the tergum. RC—rostrocarinal
width—is the width across capitulum from rostrum to carina. For convenience, 1 RC is approximately equal to 0.84 CH. PH (or PL)—height (or length) of peduncle—
is the length of the peduncle from the basal disc to the base of the capitulum.
Geographical distribution
Hoek (1883) reported that the genus Pollicipes had a wide distribution but none of the species known to him had been “found at a depth of even 10 fathoms”. He
could not neglect the possibility that future investigators might find Pollicipes in deeper water especially as species of Scalpellum were regularly brought up from the
deep sea (Hoek 1905). So far this has not been the case. The genus Pollicipes contains three of the very few intertidal pedunculates. They are found in three areas of
the world and sometimes form a most beautiful and striking component of the shore fauna. P. polymerus forms very characteristic clumps on intertidal rocks on the
Californian coast of North America (Fig. 1). A similar appearance is seen in P. pollicipes in Europe (see Figs 11 and 12 in Beauchamp 1923).
P. elegans
P. elegans is referred to by Newman & Foster (1987) and Newman (1993) as a paramphitropical species. According to Newman & Foster (1987) “the term
paramphitropical is proposed here to cover transtropical species, genera, or even higher taxa that show preferences…for the extremes of latitudinal range.” The gap in
distribution may not be complete when a genus has a species in the tropical belt but the majority of its species are north and south of it. At its northern limit P. elegans
overlaps P. polymerus.
P. elegans is found on the west coast of South America from Mexico (Hoek 1883) to Peru (Darwin 1851, Hoek 1883, Weltner 1897, Gruvel 1905, Nilsson
Cantell 1957) but does not extend as far south as Chile. In Mexico it has been recorded from Lobos Island (Darwin 1851, Gruvel 1905, Pilsbry 1909) and on an
exposed rock at Tehuantepec (Darwin 1851). In Peru it has been found at Payta (or Paita) attached to wooden posts (Darwin 1851, Gruvel 1905, Pilsbry 1909).
Bernard (1988) gives Punta Aguja in northern Peru at 6°S as the known southern limit. Some of these are obviously the same finding being recorded by different
authors when cataloguing collections in various museums. There is a puzzling record of occurrence in Java (Gruvel 1901–2, 1902, 1905) and it seems almost certain
that this is a wrong identification (Hoek 1907, NilssonCantell 1957).
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Figure 1 Pollicipes polymerus at Point Latigo, southern California showing the characteristic dense aggregations and association with
mussels.
P. polymerus
P. polymerus is one of the common animals of the intertidal region of the more exposed parts of the north American west coast (e.g. Barnes & Reese 1960, Ricketts
et al. 1968) extending from 64°N (Pilsbry 1907, Krüger 1911b) to about 27°N (e.g. Darwin 1851, Newman & Abbott 1980). In Plover Bay, Bering Strait Pilsbry
(1907) records it being associated with Conchoderma auritum and Coronula diadema on a humpback whale (Cornwall 1955b). So far Pollicipes polymerus has
not usually been regarded as a fouling organism on the hulls of ships but it has been found there (Cornwall 1955a,b). It does occasionally foul laboratory seawater
systems such as at the Scripps Institution of Oceanography, La Jolla (Newman & Abbott 1980) and at the Marine Station of the University of California at Santa
Barbara (Page 1984). The species has been frequently recorded as a member of the intertidal cirripede community from British Columbia, Washington State, Oregon
to California and as far south as Ensenada, Lower California (Hoek 1883, Weltner 1897, Gruvel 1905, Pilsbry 1907, NilssonCantell 1921, 1931, Cornwall 1925,
1951, 1955a, Shelford et al. 1935, Henry 1940b). The animal normally favours exposed positions on outer coasts (Dayton 1971) but it may occur in such areas as
Puget Sound, Washington, where the local topography produces the necessary exposure (Pilsbry 1921, Shelford & Towler 1925, Rice 1930, Towler 1930, Henry
1940a, Metz & Mueller 1975). When examining material from the Mortensen Expedition, Broch (1922) recorded what he called varieties of P. polymerus: forma
typica from La Jolla and forma echinata from Bird Rock, La Jolla and San Pedro—all in California.
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According to Tarasov & Zevina (1957, cited in Bernard 1988) P. polymerus is found on exposed rocky sites on Sakhalin Island and from the Aleutian Archipelago
to North America. Daniel (1953, 1956) found several pedunculate barnacles on a floating piece of wood on the Royapurum shore of the Bay of Bengal. This he
identified as a new subspecies Pollicipes polymerus madrasensis. The occurrence was so unexpected that Newman & Killingley (1985) decided to investigate it
further. They made an estimation of the temperature at which one of the Indian specimens must have grown by an analysis of the oxygen isotope fractionation in its shell
and compared this with the shell of a P. polymerus from California. They concluded that the Indian specimen must have developed at a temperature of about 16°C,
that is in nontropical waters. As the barnacles were found on a piece of floating wood it must have drifted into the Bay of Bengal from the colder waters of the North
Pacific.
It is of interest that the most southern records of P. polymerus at Punta Abreojos and Punta Santa Domingo on the seaward coast of Lower California (Mexico) are
at approximately the same latitude (26–27 °N) as the northernmost record, Lobos Island on the Mexican coast in the Gulf of California, for P. elegans (Newman &
Killingley 1985, Bernard 1988).
P. pollicipes
The European species, P. pollicipes appears to have a more restricted range of distribution largely because of a curtailment northwards. The species seems to be less
eurythermal and less capable of withstanding cold, temperate conditions than P. polymerus. The “thermal equivalent” of P. polymerus might have been expected to be
common on exposed coasts of southwestern Ireland and according to FischerPiette (1936) it has been found there. According to Darwin (1851) there is a record of it
being found attached to woodwork near Dublin and on a piece of drift wood in the Firth of Forth, Scotland. Darwin also records it as present on the hull of a wrecked
vessel being towed into Dartmouth, southwestern England. Newman & Killingley (1985) quote Turk (1984) regarding a record from Lands End, southwestern
England. Some of these records are very old and have not been confirmed by any later occurrences.
The species is rarely found in the English Channel (FischerPiette 1936) but is found in Europe from the north coast of Brittany in France, to Spain and Portugal, into
the Mediterranean, and in North Africa in areas of favourable exposure (Darwin 1851, Hoek 1883, Pilsbry 1907, Gruvel 1920). Joubin (1906, 1907) and Fischer
Piette (1936) recorded it on the north coast of Brittany and it is on the small islands near Roscoff (Gruvel 1902, Beauchamp 1914, Hui 1983a,b). It has been found on
the west coast of Finistère by Gruvel (1902) and FischerPiette (1955). Prenant & Teissier (1923, 1924) found it on the southwestern coast of Brittany. On the French
Atlantic coast (Bishop et al. 1957) the species is common among mussels that replace Fucus in exposed situations such as on the islands of Ouessant (Crisp &
FischerPiette 1959), Quiberon (Joubin 1906), and Yeu (GuerinGanivet 1907, Beauchamp 1923). It is on the côte Sauvage (Joubin 1906, 1907, Prenant 1932) and
1’ Archipel des Glénans (GuérinGanivet & Legendre 1909).
It is present at Biarritz and extends into Spain at St Jean de Luz and along the northern coast (Gruvel 1902, Beauchamp 1907–8, FischerPiette 1935, Navaz 1948
cited in FischerPiette & Prenant 1956) to San Sebastian, Santander and La Coruña (Bolivar 1892, Gruvel 1920, FischerPiette 1935). It does not occur in the Ria de
Vigo (Navaz 1942, cited in FischerPiette & Prenant 1956) but does occur on the west (seaward) coast of the lies de Cies at the entrance
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to the Ria (Ardré et al. 1958). It also occurs at Punta del Castro and on Cabo Silleiro at the entrance to the Ria (Barnes & Barnes 1966). On headlands on the west
coast of Portugal it is recorded by Darwin (1851), Weltner (1897), Nobré (1901), FischerPiette & Prenant (1957), FischerPiette (1959, 1963), Barnes & Barnes
(1966), and Cruz (1993, pers. comm.). Barnes & Barnes (1964) recorded the species in Catalan Bay, Gibraltar. It extends into the Mediterranean on the southern
coast of Spain and France (Hoek 1883, Gruvel 1901–2, Beauchamp 1907–8, Caziot 1921, NilssonCantell 1931). The species is also found on the African side of
the Strait of Gibraltar and on the Mediterranean coast such as at Oran and Algiers (Llabador 1937). The animals at Oran were so small that Llabador (1937) called
them var. minor. Fischer (1871) and FischerPiette (1959) record it at Cap Spartel and Arcila. On the Atlantic coast of Africa the species occurs in Morocco and
Senegal (Gruvel 1905, 1912, Bouvier 1910, Broch 1924, 1927, Burollet 1935, NilssonCantell 1939, Stubbings 1965, 1967). The most southerly record on this
coast seems to be near Dakar at about 15°N (Sourie 1954). Although well represented here, Sourie (1954) did not find it at Conakry (about 9°N).
Broch (1924) gives a record of occurrence at Teneriffe in the Canary Islands and Weltner (1897) records it in Portugal or the Azores. There is no confirmation of
this occurrence in the Azores (Stubbings 1967). Weltner (1897) also has a record of it in China, and Krüger (191 1a, b) mentions a find in East Asia but these records
seem doubtful. Krüger (1911a,b) also records it at Jan May en (71°N). This seems very unlikely unless the barnacle was attached to a whale or some other mobile
object.
Ecological habitat
P. elegans
Apart from Darwin’s (1851) statement that P. elegans occurs on an exposed rock on the west coast of Mexico nothing seems to have been published on the habitat of
this species. Much more is known about P. polymerus and P. pollicipes.
P. polymerus
It is impossible to discuss the vertical distribution of the intertidal cirripede species relative to each other on the north American coast without considering the dominating
PollicipesMytilus californianus community on these shores (Ricketts et al. 1968, Paine 1974, Seapy & Littler 1978). On the open coast in the northern part, dense
beds of mussels and their associates dominate the exposed midlittoral region. Paine (1974) found that in the PollicipesMytilus communities the Pollicipes might
disappear after about six years due, he thought, to pressure from the expanding mussel community constricting the peduncles of the Pollicipes. Barnes & Reese (1960)
thought the two species could coexist because Pollicipes could extend its peduncle and so lift the capitulum into a feeding position above the mussels. On vertical cliffs
or on overhangs on exposed coasts of Washington State P. polymerus outcompetes Mytilus sp. perhaps because the former has a stronger attachment to the
substratum, or there is a tendency for Mytilus to prefer lower levels (Paine 1974).
In the most generalized situation the PollicipesMytilus community is interposed between Chthamalus dalli and Balanus glandula above and Semibalanus
cariosus below it (Dayton
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1971). Depending on local conditions the situation becomes modified and the separation less marked. With increased shelter the PollicipesMytilus community is
absent and the general zonation is then Chthamalus dalli, Balanus glandula, and Semibalanus cariosus. On the southern part of the coast in exposed areas the
PollicipesMytilus community separates Chthamalus fissus and Balanus glandula from Tetraclita squamosa rubescens (Hewatt 1935, 1937, Shelford et al. 1935,
Barnes 1960).
It is of interest to record the ecological succession on an artificially cleaned area (lyd square) in the mussel zone at Cabrillo Point, Monterey Bay (Hewett 1935). The
area was protected from the direct force of the waves but was subjected to strong currents of welloxygenated water. The tidal level at the top of the cleaned area was
4.8ft. in a tidal range of 2–7ft. Prior to cleaning, the area supported a typical Mytilus association. In the middle of the lower part of the area there were 356 Pollicipes
polymerus and 1612 Mytilus californianus on 23 November. After cleaning, the area was examined at intervals and the results for these two species are given in
Table 1. Pollicipes was recorded before any Mytilus; it is not clear whether the former were actually attached to the rocky substratum or to some other organisms that
were already present by then. The Pollicipes were said to be 14mm (but what measurement was made is not recorded) on 4 March, so cyprids must have settled in
February or earlier; they may have been too small to notice on 19 February. Initial growth must have been fast as has been noticed in other habitats (see Table 15, p.
359). Hewatt (1935) offered no explanation for the decrease in number of Mytilus recorded on 18 July. Once established at 109 by 25 April Pollicipes polymerus
remained stable suggesting that there was no settlement after March. This is curious as according to Hilgard (1960) the breeding season at Monterey Bay is April to
December, with peaks in June, September, and December. Even after 2½yr the “cleaned” area had not reverted completely to the original cover before cleaning took
place.
Table 1 Ecological succession of Mytilus californianus and Pollicipes polymerus on a cleared area at Cabrillo Point, MontereyBay. (After Hewatt
1935).
Mytilus Pollicipes
Date Number Size (mm) Number Size (mm)
23 November before cleaning 1612 – 356 –
19 February – – – –
4 March – – 76 14
25 April 55 4 109 20
3 June 51 11 109 24
18 July 7 – 109 –
13 October 53 – 109 –
26 December 89 – 108 –
Away from the open coast, where both the dominating Pollicipes—Mytilus association and Tetraclita squamosa rubescens are absent, the whole intertidal is open to
competition between Chthamalus fissus and Balanus glandula. As conditions become less favourable to the latter species, in particular in higher temperatures,
Chthamalus fissus may virtually cover the whole midlittoral. At Ensenada in Lower California only isolated individuals of Balanus glandula and Tetraclita squamosa
rubescens may be present with small groups of Pollicipes polymerus in the most exposed places.
On the coast of Oregon, Giesel (1969, 1970) records an association between Acmaea digitalis and Pollicipes polymerus. The limpet may be found on sandstone
faces in the high intertidal or on the valves of P. polymerus (usually the scutum). This association influences
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the growth form and shell pattern of the limpets living on the barnacle compared with those living on the rock surface.
P. pollicipes
P. pollicipes is present on the northern coast of Brittany on the most exposed cliffs and reefs (Joubin 1906, FischerPiette 1936). Joubin (1906, 1907), Guérin
Ganivet & Legendre (1909) and Beauchamp (1914) associated the species with deep channels in steep cliffs characteristic of the wild areas of the western coast of
Brittany. It is also found in caves at the ends of these channels where the waves break with extreme violence. Joubin concluded that the species lived in water that was
constantly agitated and oxygenated. The channels may be 100m deep and the Pollicipes formed a layer above the Fucus zone. When the sea was too rough mussels
replaced the Fucus. This mussel bank was interspersed with groups of 30–40 individuals of Pollicipes pollicipes. These groups were firmly attached to the rock and
to one another and could not be easily removed (FischerPiette 1936). The frequent association of mussels and Pollicipes on the Atlantic coast of France was also
noted by Beauchamp (1923) and Prenant (1932). The species was associated with mussels on the many small islands in TArchipel des Glénans and formed a marked
zone between the upper Chthamalus stellatus and the laminarians (Prenant 1932, Bishop et al. 1957, Crisp & FischerPiette 1959). Balanus perforates was found
below the Pollicipes in some places (Prenant 1932). At Biarritz, FischerPiette (1935) mentions P. pollicipes as occurring below Chthamalus stellatus and Mytilus
edulis.
The basque coast is favourable to Pollicipes pollicipes, so much so that it was found not only on the exposed coast but in some rias such as Ria de Pasajes (Bolivar
1892, Navaz 1948 cited in FischerPiette & Prenant 1956, FischerPiette 1955, Bishop et al. 1957, Crisp & FischerPiette 1959). It was not, however, found in the
Ria de Vigo in Spain (Navaz 1942 cited in FischerPiette & Prenant 1956, FischerPiette 1955) but it has been found on headlands at the entrance to the Ria (Barnes
& Barnes 1966) and in areas facing the open sea (Ardré et al. 1958). At Cabo Silleiro the Pollicipes were embedded in rock fissures and Chthamalus stellatus was
found on the open rock surfaces. Here Balanus perforates was restricted to gullies on the landward side of the Cape and at Punta del Castro it was restricted to caves
(Barnes & Barnes 1966).
On the Portuguese coast Pollicipes pollicipes was found in crevices and places where there is a lot of surf and strong undertows (FischerPiette & Prenant 1957).
It was also connected with strong surf on the Atlantic coast of north Africa and with Mytilus africanus at Cap Spartel (FischerPiette & Prenant 1957). At Arcila,
FischerPiette (1959) recorded rare and large Pollicipes in crevices of cliffs facing the sea. Groups of mussels and seaweeds, Corallina and Codium, were also
present. At Cap Blanc, Pollicipes were again common in rock fissures at the northern boundary of the Bay of Dakar (Gruvel 1912). Many intertidal invertebrates on
the coast of Senegal are protected from extreme desiccation by their walls retaining a certain amount of water (Sourie 1954). P. pollicipes is one of the examples
given. Fissures and crevices are the favourite habitat and this must provide some extra protection for the stalks of these animals. Only the capitulum appears above
these crevices. In this region of the Atlantic coast of north Africa chthamalids are found above Pollicipes and Balanus tintinnabulum below. Stubbings (1965) in the
Dakar region, found Pollicipes pollicipes with young attached and also epizoic Chthamalus dentatus. He also found a group of 50 or more Pollicipes growing close
to Balanus tintinnabulum.
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Description of adults
The terms used to describe parts of the pedunculate cirripedes were clearly defined by Darwin (1851, pp. 3–7) and many of these are still in use today. Essentially
pedunculates are cirripedes having a flexible stalk (the peduncle) that supports the capitulum. The peduncle is muscular and contains the ovarian tissue. The animal’s
body is in the capitulum within what Darwin called “the sack” but what is now usually referred to as the mantle cavity. The body consists of a thorax from which the cirri
arise. These have two arms, the rami, supported by a common segment, a pedicel. The prosoma is an extension of the thorax and contains the stomach. Two small
projections that may be uni or multiarticulate are on each side of the anus, and just above the proboscislike penis are the caudal appendages. The base of the penis is
also called a pedicel. On the thorax, prosoma, and pedicels of the cirri there may be filamentary appendages. The mouth consists of the labrum with mandibles,
maxillae, and outer maxillae.
The number of plates and valves protecting the capitulum can vary from 18 to more than 100 (Fig. 2). This number can vary considerably in the same species and
generally increases with age. The number of whorls of plates at the base of the capitulum is important and also increases with age, although there are conflicting reports
of this, in the three Pollicipes species. The arrangement of plates on the capitulum of P. elegans and P. pollicipes as well as a schematic drawing of the former species
are shown in Figure 3. P. elegans is regarded as the most primitive of the three Pollicipes species because the rostrolatus and carinolatus are the least separated from
the median latus (Newman 1987). The rostrum is small and rather unimportant in all three species.
The length of the peduncle may vary according to species and environmental conditions. In each of the three species it is covered with calcified scales placed
alternately and symmetrically. The size varies with the age of the individual; the lower scales, formed when the animal was younger, are smaller than the upper scales.
The spaces between the scales are
Figure 2 Pollicipes pollicipes (A) and P. polymerus (B) as illustrated by Darwin (1851)
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Figure 3 Pollicipes elegans: schematic drawing (A) and diagramatic arrangement of capitular plates (B) compared with P. pollicipes
(C). C=carina; c=imbricating plates added between L and C; CL=carinolatus; L=lateral or median latus; R=rostrum;
r=imbricating plates added between R and L; RL=rostrolatus; S=scutum; sr, 1 and sc=imbricating plates below R, L and C,
respectively; T=tergum; ••••= small plates added from below. (After Newman 1987).
greater nearer the base of the peduncle owing to stretching and contracting and renewal of intervening membrane. The scales themselves are covered with a membrane
as are the valves on the capitulum. Sometimes this membrane may be abraded.
Darwin (1851) gave a detailed description of each of the three species of Pollicipes so only the essential points are given in Table 2. Occasionally the descriptions
have been augmented by descriptions published after Darwin (1851). All three species are hermaphrodites. Gruvel (1905) and NilssonCantell (1921) stated that the
scales on the lower part of the peduncle of P. polymerus were in the form of spines irregularly arranged; Broch (1922) and Henry (1940b) did not agree. Two forms
of P. polymerus, forma echinata and forma typica,
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Table 2 Description of various parts of the three Pollicipes species; details taken mainly from descriptions by Darwin (1851) augmented by those of
Gruvel (1902, 1905), Pilsbry (1909), Krüger (1911a,b), NilssonCantell (1921), Cornwall (1925, 1951, 1955a,b), Broch (1927) and Henry (1940b).
Approximate dimensions of adult taken from Gruvel (1905); length, tip of tergum to junction with peduncle; breadth, widest part.
P. elegans P. pollicipes P. polymerus

Capitulum More globular than P. pollicipes; 1 to 3 whorls of Almost triangular, occluded edge tinted red; 1 Triangular, slightly compressed, broad with scuta and terga
plates below rostrum, reddishorange colour with to 3 whorls of plates below rostrum, whitish at an oblique angle with respect to peduncle; 2,3 or more
parallel markings not seen in P. pollicipes; scales grey colour; scales flattened, close together, whorls of plates below rostrum, whitishgrey colour; scales
flattened, close together, regular and overlapping, regular and overlapping, ≤0.5mm long. in upper whorls flattened, close together and regular, grey
redorange colour, little longer than in P. colour, very small, become progressively smaller from
pollicipes superior to inferior.
Peduncle Longer and thinner than in P. pollicipes; scales Narrow, a little longer than capitulum; small Upper part less diameter than capitulum, 2 to 3 times as
arranged in close whorls, a little larger than in P. scales arrange symmetrically in close whorls; long as capitulum; scales minute and flattened,
pollicipes, reddishorange colour. scales taper towards tip. symmetrically arranged in close whorls; scales uniform
throughout length, not tapered
Scutum Similar to P. pollicipes Oval, basal and tergolateral margins merging Irregularly oval, basal margin appears to have three parts
into each other, apex pointed; deep internal pit corresponding with margins of rostrum, rostral and adjoining
for adductor muscle. latus; tergolateral margin curved and protuberant, umbo
projects a little over scutal margin of terga.
Tergum Similar to P. pollicipes Larger than scutum, carinal margin very Broadly oval, projecting slightly beyond other valves, apex
curved and protuberant, basal angle blunt, rounded, blunt, basal angle bluntly pointed, scutal margin
scutal margin curved with upper part straight curved, occludent margin twosided at 90° to each other,
or formed by two lines corresponding to tergal length and area about equal to scutum.
margin of scutum and one side of upper latus.
Carina Curved, basal internal margin rounded not square Very curved extending up between terga, Triangular, rather narrow internally deeply concave, curves
as is usually case in P. pollicipes. widens from bottom to top, internally deeply upwards to apex, not projecting, basal margin protuberant,
concave basal margin very protuberant, basal
internal margin squarish, apex rounded.
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Rostrum Similar to P. pollicipes(?) Less than onethird length of carina, concave, triangular, Onethird of carina, triangular, nearly same as shape as
basal margin slightly protuberant. carina, internally very slightly concave, basal margn
truncated or angular in middle.
Latera Upper pair of latera (viewed Upper pair of latera, corresponding to spaces between Upper pair of latera between scuta and terga only slightly
internally) have an area about twice scuta and terga is the largest, but is almost the same size longer than latera immediately below and these are only
that of latera that corresponds to as the pair between terga and carina, next largest pair slightly larger than those below them. Valves on lowest
space between carina and terga. (rostral) is that between scuta and rostrum; some of whorl are elongated transversely and are minute, but still
lower latera are of nearly the same size. about twice the size of peduncular scales and a different
shape.
Ovigerous Consist of long prominent folds None None
fraena thinning to nothing towards bases of
scuta, no gland seen, not functional.
Caudal Multiarticulate, shorter than lower Multiarticulate, longer than lower segments of pedicel Uniarticulate, very small, blunt and rounded, few very short,
appendages segments of pedicel of 6th cirri, of 6th cirri, cylindrical, blunt ended, short row of spines thick spines at tips,
four articulations constricted near round upper edge of each segment, five oblique,
base. imperfect articulations.
Filamentary On prosoma—12 in each of 4 On prosoma, 13–14 pairs, short, blunt filaments in 2 On prosoma—12–14 pairs in 2 rows, middle filaments
appendages longitudinal rows, filaments in 2 longitudinal rows; those nearest thorax are longest. longest equal to about half diameter of thorax, each is
outer rows half length of those in On base of 1st cirrus—a pair, shorter and with a few flattened and tapered a little towards the tip, which has
inner 2 rows; those nearest thorax minute’ spines. serrated crests on both sides, directed downwards towards
are longest. On mouth—pair each side, pointing upwards, thinly each other and the thorax.
On base of 1st cirrus—a pair of clothed in long spines. Testes in all filaments. On base of 1st cirrus—on each side a short appendage with
filaments pointing upwards half a few bristles at end.
length of these on prosoma. On mouth—at each side a long, dark coloured appendage,
On mouth—thicker filament each with a few scattered bristles, directed upwards towards
side, pointing upwards, minute adductor scutorum muscle.
scattered spines. Some apices have
minute pectinated scales. Testes in
all filaments.
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Labrum Similar to that of P. pollicipes. Highly bullate, longitudinal diameter equal to rest of Highly bullate, upper part less so than lower part, small
mouth, upper part square not overhanging lower part, blunt teeth and some bristles on crest.
small teeth on crest.
Palpus ? Oval, outer and inner margins almost alike, thickly Clubshaped, approaching but not touching, broad or
clothed with spines square extremities, thickly fringed with serrated bristles.
Mandibles Interspace between third tooth and inferior Three strong yellow teeth, inferior point broad and Three very strong teeth, slightly graduated in size,
angle pectinated. coarsely pectinated. inferior angle very coarsely pectinated, lower edges of
main teeth roughened.
Maxillae Similar to P. pollicipes but has 6–8 pairs of Three tufts of fine bristles separated by large spines, Spinose edge about half length of mandibles, two upper
spines between the two upper tufts of fine first tuft near to upper spines, second tuft in middle, spines very strong almost covering a tuft of fine spines,
spines. third tuft at inferior angle, second and third tufts stand in the middle there is a second tuft of spines and at the
on prominences, between first and second tufts are 3 inferior angle a third—both on prominences; in
pairs of strong spines and between second and third interspaces between tufts are 3 or 4 pairs of spines,
are 4 more pairs.
Outer Similar to P. pollicipes Inner edge has a central notch with bristles above and On inner margin a deep notch with a compact row of
maxillae below marking two equal combs; on exterior surface serrated bristles above and below it.
bristles are longer and more spread out.
Cirri 1st—more than half number of segments of 1st—short and thick, less than half number of 1st—same number of segments in each ramus, with
6th cirrus, lower segments thick with doubly segments in shorter ramus than in 6th cirrus, 1st pair many bristles and tufts of fine spines, near to 2nd pair of
serrated spines; lower segments of anterior not far removed from 2nd pair, lower segments of 1st cirri with twice as many segments in rami; among the
rami have spines; these spines have peculiar cirri, thick, protuberant with doubly serrated spines; ordinary spines there are very strong ones with doubly
bent teeth (not seen in P. pollicipes); 1st lower segments of 2nd cirri protuberant with non pectinated teeth, many spines are doubly serrated;
and 2nd pairs of cirri have more segments serrated spines; lower segments of anterior ramus of posterior cirri moderately long, curled, segments
in proportion to the 6th pair than in P. 3rd cirrus protuberant; posterior rami of first 3 pairs flattened, anterior surface protuberant with 6 spines,
pollicipes. only slightly protuberant and resemble 3 between spines are large tufts of fine spines.
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P. P. pollicipes P. polymerus
elegans
Cirri posterior pairs of cirri, segments of these pairs have 6 pairs of short, strong spines, with thick brush of short fine
(continued) bristles in between, thick minute spines on upper lateral edges of segments.
Penis ? Purple, short, stout, with fine spines Dark coloured, short, stout,
with fine spines.
Capitulum
Length 21 mm 18 mm 18 mm
Breadth 18 mm 19 mm 16 mm
Peduncle
Length 50 mm 30 mm 35 mm
Diameter 13mm 11 mm 10 mm
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were recorded in southern California by Broch (1922) but they have rarely been mentioned by other workers.
Fracturing of lamina
In P. polymerus the outer lamina of one or more of the capitular plates may split (Cornwall 1925). Sometimes when small balanid barnacles are growing on the plates
the outer laminae are quite loose and about half the thickness of the plate eventually breaks off. When the scutum is split in this way the depression for the attachment of
the adductor muscle shows on the outside as a prominence. This splitting of the valves is in no way analogous to moulting of the soft parts. With very few exceptions the
calcareous covering is not moulted or cast (but see p. 355). Cornwall (1925) examined capitular plates of P. polymerus on which Semibalanus cariosus had settled.
This latter barnacle has a membranous base and the bases of the wall plates firmly adhere to the capitular valve of the pedunculate. During growth of the balanid the
increasing circumference of the shell sets up a tension on the valve that eventually causes it to split off the lamina. Cornwall (1925) found all stages of this process in a
collection of Pollicipes polymerus from Departure Bay, British Columbia. A similar phenomenon has been reported for P. pollicipes (T.Cruz pers. comm.) from the
southwestern coast of Portugal.
Abnormal shell form

It has been shown (p. 310) that Pollicipes species favour habitats with strong wave action such as deep gullies where the backwash of sea water rushes down. P.
polymerus is associated with high energy intertidal areas along the Californian coast, USA (Ricketts et al. 1968). During a biological survey of caves on Southeast
Farallon Island, California, several groups of P. polymerus were found in a moderatelyexposed dimlyilluminated side channel within a cave (Chaffee & Lindberg
1980). They found, respectively, two and three atypical individuals centrally placed in two groups of about 20 typical P. polymerus. An atypical animal had a carina
that extended up to 1 cm beyond the apex of the terga, a reduced rostrum, the terga and scuta showed signs of exfoliation (see above), and accessory plates and
peduncular scales were subrectangular in shape. The membrane surrounding the aperture was red, an indication of reduced light conditions. The authors proved
statistically that the capitular growth of the atypical animals was allometrically different from typical ones. Allometric growth may result from competition in a crowded
group of barnacles although it is difficult to see what advantage could be gained from a projecting carina.
Integument and calcified parts
Outer integument
The outer integument of a pedunculate cirripede forms the outer covering of the peduncle and the capitulum. It is thick and laminated (Thomas 1944) and incorporates
a variable number of calcareous plates on the capitulum and numerous calcareous scales on the pedun
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cle (Mahmoud 1959a). A thick protective covering, which is not lost during moulting and has the ability to produce calcareous structures, is an advantage in a sessile
animal that has no escape from predators or changing conditions of the environment. The structure of the integument, particularly in P. pollicipes, was examined
histologically over 100 years ago by Koehler (1888, 1889a,b) and by Gruvel (1904a, 1905). Later Prenant (1924) thought some of the older work needed
clarification. Koehler (1889a) found the epithelium covering the outside of the peduncle in P. pollicipes to consist of a layer of cylindrical cells placed regularly side by
side. They had an oval nucleus and normally contained pigmented granules. The epithelial layer, therefore, formed a black border of 30μm thickness which extended
for the whole length of the peduncle. The outer covering consists of a thin outer epicuticle that is homogenous, refractile and ambercoloured and an inner endocuticle of
chitin that is laminated and colourless (Thomas 1944, Mahmoud 1959a).
In Pollicipes the plates and scales project through the endocuticle. Tegumental glands lie immediately below the epithelium (Thomas 1944, Mahmoud 1957/58) and
ducts pass through the chiton of the endocuticle (Darwin 1851). Koehler (1889a) described canals (ducts) in the integument of P. pollicipes some of which went to the
calcareous scales on the peduncle. At the base of the scale the duct formed a swelling (the socalled “organes de Koehler”) from which fine ducts penetrated through
the scale to the surface of the peduncle. The formation of the scales has been described by Koehler (1889b) and Mahmoud (1959b) and figures illustrating this have
been given by Broch (1922), Prenant (1924), and Mahmoud (1959b).
The thick cuticular layer covering the peduncle of Pollicipes is covered with numerous semiglobular elevations. These are arranged symmetrically and alternately in
close whorls round the peduncle. There are small calcified scales embedded in these elevations (Mahmoud 1959a). In P. pollicipes the scales are pearshaped with the
rounded edge embedded; they curve and project upwards. The scales are formed of calcite crystals arranged at 90° to the ectoderm and they are covered with a
brown cuticular sheath. Fully grown scales have an organic matrix, as seen when decalcified. A fine, ambercoloured, chitinous duct (the tegumental gland duct, passes
from the ectoderm, through the basal wall of the scale and on to its outer surface. In scales in the upper whorls there may be two or three separate ducts. Longitudinal
sections of the peduncle indicate that lower down the peduncle the scales become smaller. Damage caused, for example, by abrasion can be repaired by secretion from
the gland cells via the tegumental gland ducts to the outer surface of the scales.
Adults
Gruvel (1899) admitted the inconsistency in number and position of capitular plates on pedunculates. This applied not only to different species but also to members of
the same species (Darwin 1851, Gruvel 1905, Mahmoud 1959a). Gruvel (1899) remarked that capitular scales were only peduncular scales modified and adapted to
the special function of protecting the prosoma and he was led to admit that Darwin’s (1851) separation of peduncle and capitulum was arbitrary. Joleaud (1910a,b,
1916) mentioned the evolution of primitive calcified plates on the capitulum of cirripedes. The general arrangement of plates in P. pollicipes was discussed by him
(Joleaud 1913, 1916). He gives an arrangement in two cases; one animal had 31 plates and 43 small pieces (his words) in five rings and the other had 39 plates and 62
small pieces in six rings. These animals were the same species but different ages. Joleaud considered this plan of plates instructive in a study of the development of the
capitulum in Pollicipes. Nussbaum (1890) mentioned the growth of the capitular plates and
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examined some of the glands histologically in P. polymerus. Broch (1922), however, felt that Nussbaum’s figures were not very clear.
Broch (1912) followed plate development in pedunculate cirripedes and later published a preliminary report (Broch 1921), and then a full account (Broch 1922) of
the growth zone and the structure of the capitular plates and peduncular scales in P. polymerus. The growth of the animal is restricted to the transition zone between
the capitulum and peduncle (Mahmoud 1959b). Krüger (1940) mentions the plate development in both P. polymerus and P. pollicipes.
In P. pollicipes the capitular plates are calcified layers of fine evenlydistributed calcite crystals. The crystals are arranged at 90° to the basal epithelial layer of the
plate. The calcified layers are separated by chitinous sheets (Gruvel 1905, Broch 1922) as seen after decalcification of the plates. The term “growth lines”, applied to
these chitinous sheets by Broch (1922), is misleading according to Mahmoud (1959a). Fine ambercoloured tegumental gland ducts run through the different laminae of
the plates. There is a thick brownish chitinous sheath covering the plate externally. In P. pollicipes sections show that the number of laminae in each layer is variable
even in the same plate as well as in different plates of the same animal (Mahmoud 1959a). The observations of Mahmoud on “lines of growth” agree with those of
Broch (1922) on P. polymerus but in neither case can they be used to determine the age of the animals.
Juveniles
Broch (1922, 1927) found cyprids crawling on the capitulum of adult Pollicipes but settlement and developing young were only found on the peduncle. He describes in
detail the development of the plates and scales of juvenile P. polymerus (Broch 1922). As soon as the cyprid settles and is attached to the substratum, five primordial
valves appear—the embryonic carina, paired terga, and paired scuta. These are chitinous, have a porous nature and contain no calcium carbonate. Their calcification
begins on the interior giving them the appearance of a scale indicating the umbo of the plate. The umbones are apical so calcification continues only along the lower
sides of the plates. The rostrum appears as soon as calcareous deposits become evident outside the margins of the primordial valves. The rostrum is soon followed by
an upper latus below the interval between the tergum and scutum. The next plates to be distinguished are a carinal and a rostral latus and at the same time the first plate
of the lower row of latera appears below the interval between the rostrum and rostral latus (Fig. 4). The following latera of the lower row quickly appear as well as the
subcarina.
The peduncular scales do not appear until the subcarina has developed. The scales on the rostral side develop first followed by the lateral scales nearer the carino
sagittal line. Owing to the growth of the peduncle, which is limited to the “growth zone” (Mahmoud 1959b), the peduncular scales form an oblique series ascending
from the rostral to the carinal side of the peduncle. Peduncular scales produced in this region are gradually pushed lower down the peduncle by the production of
younger scales above them (see Fig. 14, p. 354). Gradually lower rows of latera develop on the capitulum, with the new plates below the intervals in the preceding
row. The growth of the capitulum eventually becomes very slow (see Table 15, p. 359) and the formation of new latera may cease. Growth of the peduncle and the
production of peduncular scales appears to continue throughout the life of the individual and always in the same way. Thus, the oblique serial arrangement of scales is
maintained although in older animals it may be difficult to see due to damage or accidental contractions of the peduncle.
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Figure 4 Early plate development in Pollicipes pollicipes (A) and P. polymerus (B). 1–4=advancing stages of growth; C=carina;
CL=carinolatus; L=median latus; P=peduncle; R=rostrum; RL=rostrolatus; S=scutum; SL=sublatus; T =tergum; not to scale
(modified from Broch 1922, 1927 and Anderson 1994).
The development of the skeleton of juvenile P. pollicipes follows a similar pattern and Broch (1927) has described it for animals found on the Atlantic coast of
Morocco. As in P. polymerus the first latus does not appear until the rostrum is comparatively large. The scales on the peduncle begin to appear as the subcarina
develops. The latera for the capitulum and the peduncular scales are only formed in the “growth zone” between the capitulum and the peduncle. The peduncular scales
are continually produced as in P. polymerus and appear to stop development soon after they leave the growth area.
Inner integument
The inner integument of a cirripede is the thin covering of the mantle cavity and the soft parts of the body and it is shed regularly during moulting (Thomas 1944).
According the Koehler (1889b) this internal “epithelium” is usually devoid of pigment in pedunculates but in Pollicipes there is some pigmentation although not as
intense as that found in the external “epithelium”. The glands renewing the layer shed during moulting are active only during moulting. These are found near the maxillae,
which prompted Gruvel (1940b) to call them “salivary glands”. Thomas (1944) regarded this as misleading and preferred labial or suboesophageal glands. They may
be connected with the production of cuticle during moulting. The cuticle is thickest in the vicinity of the gland cells but becomes thinner as the distance
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from them increases. Barnes & Reese (1959) refer to glands connected with feeding on the labrum and outer maxillae as well as the tegumental suboesophageal glands
near the bases of the first three pairs of cirri.
In pedunculates all the gland cells regenerate after secretion. The cement glands (see below) are always active in order to maintain the adhesion of the peduncle to
the substratum. The glands in the outer integument supplying the scales may be active at any time to repair damage and produce new scales. The glands supplying the
inner integument only secrete during moulting.
Adhesion and cement

One of the necessities of intertidal cirripedes is their ability to attach themselves to the substratum. This adhesion must be strong enough to hold the animals in place
despite repeated battering by strong seas in exposed positions, the whiplash effects of seaweed, attacks by predators and even attempts by man to dislodge them
(Bernard 1988). The production of the adhesive fluid involves exocrine secretions (Anderson 1994). The mechanism for producing the fluid and its passage to the base
of the peduncle in the case of stalked barnacles have been the subjects of many investigations.
Cement glands were mentioned by early workers such as Darwin (1851, 1854), Krohn (1859), Koehler (1888, 1889a), Nussbaum (1890), and Gruvel (1904a,
1905). Until comparatively recently there had been little further work on the specialized glands responsible for the secretion of cement and that was in balanomorphs
(Yule & Walker 1987, Anderson 1994). Pedunculate barnacles, especially Lepas species were considered by Lacombe & Liguori (1969). The adhesion mechanism in
adult pedunculates consisted of secretory cells intermingled with the ovarian tissue and linked by a duct system leading to an opening through each antennule at the base
of the peduncle in Lepas species (Lacombe & Liguori 1969, Lacombe 1970). These authors also implied that the structure of the cement glands might be less complex
in lepadids than in balanomorphs. In Lepas fascicularis large amounts of cement are discharged around the base of the peduncle. In this species the cyprid usually
settles on a relatively small and floating object. The mass of secretion at the base of the peduncle of the young adult increases with age and acts as a float (Barnes &
Blackstock 1974).
The properties of the cement, its biochemical composition and its tensile strength have been thoroughly examined in balanomorphs (see review by Yule & Walker
1987). The cement produced by pedunculate barnacles has not received so much attention (Barnes & Blackstock 1974, 1976, Walker & Youngson 1975) and this
work has all been on Lepas species.
Darwin (1851) reported that in Pollicipes polymerus two cement ducts ran down the peduncle within the longitudinal muscles and close to the ovarian tubes. The
ducts penetrated the base of the peduncle and “cement poured forth”. The cement glands were found near each other high up in the peduncle. He did not mention the
attachment or cement apparatus in P. pollicipes or P. elegans. Koehler (1888) examined Pollicipes (probably P. pollicipes) but the species was not given. He found
the cement gland in the basal portion of the peduncle.
The cement cells and ducts as well as the properties of the cement have apparently never been examined recently in Pollicipes species, but they may be expected to
be similar to those in Lepas species. There appears to be only a personal observation of G.Walker and mentioned in Naldrett’s paper (1993) that large quantities of
the liquid cement from Pollicipes species turned opaque in sea water. This apparently agrees with Naldrett’s (1993) observation that liquid
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cement does not solidify if diluted sufficiently with a tris buffer containing no salt. The cement must remain fluid while in the cement ducts and Naldrett suggested that the
cement is secreted in a low salt or saltfree solution which flushes the cement through the ducts. Solidification of the cement would then only occur on contact with salt
water outside the duct system.
Muscles
The musculature of cirripedes is elaborate and involves the control of various parts of the animal. In a pedunculate such as Pollicipes the largest muscles are in the
peduncle and the wall of the capitulum; they are also the simplest. There are more complicated small arrangements of muscles in the prosoma and thorax including the
cirri, penis, and mouthparts (Anderson 1994). The variation of haemolymph pressure is a major factor in the movement of Pollicipes and so the structure and function
of the haemocoelic system are closely connected with those of the muscles.
Several accounts of cirripede muscles including those of Pollicipes were published in the late 1800s and early 1900s (such as Darwin 1851, Koehler 1888,
1889a,b, Gruvel 1904a, 1905). Koehler (1889a) found anastomosing fibrils and granulations in the Pollicipes peduncle but not in other pedunculates. Both Koehler
and Gruvel noted that the peduncular muscles were not striated. Some of the early papers were merely descriptions while others were supported by histological
sections. All three Pollicipes species were mentioned by Gruvel (1904a). Joleaud (1911, 1916) suggested the position of the insertion of the adductor muscle on the
scutum as a means of classification of the pedunculates. Castaño (1954, 1955) examined the muscular insertions and muscle tissue of P. pollicipes and made
comparisons with some other cirripede genera. Muscles and their ramifications in the peduncle of P. pollicipes have been described by Garrault (1934). She found that
the loose connective tissue was supported by elastic fibres forming a framework. The fibres were in direct contact with the external epidermal cells and perhaps were
secreted by them. The muscles involved in the oral cone and the mouthparts of P. pollicipes have been described in detail by Petriconi (1969).
The musculature of the pedunculate Lithotrya has been carefully described by Cannon (1947) and he pointed out where it differed from that of other stalked
barnacles including Pollicipes. He hesitated to name the various muscles merely identifying them by numbers. The most recent work on muscles of a stalked barnacle is
included in KlepaPs (1985) paper on Ibla cumingi. She describes the muscles in meticulous detail making some comparisons with Pollicipes polymerus as given by
Nussbaum (1890) and Lithotrya as given by Cannon (1947). In her table (Klepal 1985, p.71) she has named the muscles including those of Pollicipes polymerus
following the muscle nomenclature of Nussbaum (1890).
As muscles exert control over the various bodily functions in Pollicipes species more about their action will be given in the appropriate sections of this review.
The nervous system

The application of electrical recording techniques to work on the giant muscle fibres of the large Balanus nubilus and the photoreceptors of B. eburneus was the
beginning of “modern” cirripede neurobiology according to Gwilliam (1987). There were, however, much earlier
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investigations by, for example, Brandt (1871), Koehler (1889c), Nussbaum (1890), and Gruvel (1903, 1904a,b). Some of this work was done on pedunculates.
Brandt (1871) made rather general statements implying that the ganglia in pedunculates were always regularly placed behind each other and that the connectives
were of equal length. Koehler (1889c) found that this varied between species. He found that in Pollicipes the connectives between the first and second ganglia were
very much shorter than those connecting the second to the third and the third to the fourth. The fourth and fifth ganglia were so close together that they almost touched.
All these authors agreed that there were six pairs of ganglia. Gruvel (1904a) found similar variable lengths of connectives in P. polymerus. He also discussed “l’organe
de Koehler” calling it a nervous organ situated at the base of the peduncular scales in Pollicipes and apparently only found in that genus. In P. pollicipes it was massive
and multicellular, the ganglion cells being more or less swamped in a black pigment. In P. elegans the cells were dissociated and only united by prolongations that
allowed spacious intercalary spaces. The pigment too was absent, which facilitated histological examination. Thomas (1944) disagreed saying that the organs were non
cellular and merely swellings on “canals” that go through the integument to the surface. The peripheral cells usually formed at the end of the prolongations that passed in
fine channels through the scales to the surface. According to Gruvel (1903), these nervous organs probably served to inform the animal of physical changes in the
environment.
Nussbaum (1890) described in detail the nervous system including the nerve endings in P. polymerus. He found the same number of ganglia as seen earlier by
Brandt (1871) and Koehler (1889c) in P. pollicipes but he did not mention their work or make any comparisons with it much to the chagrin of Koehler (1891) and
Gruvel (1904a). Whether this was intentional or merely that he was unaware of their papers will never be known. Nussbaum included some dimensions for the ganglia
and the length of commissures between them but, unfortunately, he did not give any size for the adult cirripede in which they were measured (Table 3). These
differences in length of commissures agree with Koehler’s (1889c) finding in P. pollicipes. Nussbaum (1890) mentioned two enigmatic organs—“undefinirbare
Organe”. He thought one of these had a nervous function but Gruvel (1904a,b) decided that they were both granular.
Cornwall (1936, 1953) also described six pairs of ganglia in P. polymerus, five he called infraoesophageal and one supraoesophageal (Fig. 5). The first pair of
infraoesophageal ganglia were fused and are referred to as the suboesophageal ganglion by Batham (1945, in P. spinosus, now Calantica spinosa), by Klepal
(1985, in Ibla cumingi), and by Anderson (1994). From this ganglion the splanchnic nerves go deeply into the body and serve the vis
Table 3 P. polymerus: size of ganglia and distances between them (µ.m). (From Nussbaum 1890, Cornwall 1953).
Ganglion Size Distance to next ganglion

(Nussbaum) (Nussbaum) (Cornwall)
Infraoesophageal 700
100 Very short, hardly visible
First thoracic 200
800 Clearly visible
Second thoracic 250
1000 Clearly visible
Third thoracic 300
150 Not visible
Fourth thoracic 500
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Figure 5 Pollicipes polymerus: schematic representation of the nervous system. IVI=nerves leading to cirri; 1=infraoesophageal
ganglion lying behind oesophagus (OE); 2=supraoesophageal ganglion lying in front of oesophagus; 3=nerves connecting 1 and
2, the circumoesophageal chord; 4=nerves leading to network of nerves near body surface; 5=optic nerve; 6=splanchnic
nerves running into body; 7=nerves leading to penis. (After Cornwall 1953).
cera including the mouthparts (Batham 1945, Klepal 1985). It seems that Cornwall (1953) was wrong in saying that nerves for the mouthparts originated in the supra
oesophageal ganglion in Pollicipes polymerus. The circumoesophageal cord also arises from the suboesophageal ganglion. The next four pairs of ganglia, now called
thoracic ganglia (Barnes & Gonor 1958b, Anderson 1994) are variously spaced from each other. The distance between the suboesophageal ganglion and the first pair
is very short and hardly visible. This pair and the next two pairs of thoracic ganglia are not fused and the connectives between them can be seen. The connective
between the last of these and the final pair, which are fused together, is not discernible. Nussbaum (1890) was, however, apparently able to measure these distances
(Table 3). Nerves from the thoracic ganglia lead to the respective cirri. They divide in the pedicel, one branch going to each ramus. The third and fourth pair of thoracic
ganglia, that is, the fourth and fifth infraoesophageal ganglia of Cornwall (1953), being fused together, serve the fourth, fifth and sixth pairs of cirri. A pair of nerves to
the penis branches off the bases of the nerves to the sixth cirri.
Several small nerves pass from the supraoesophageal ganglion to the eyes and a large nerve becomes the antennary or peduncular nerve. Nerves from these ganglia
eventually form the network of nerves near the body surface (Cornwall 1953).
The nervous system of cirripedes is simpler than that of other crustaceans; there is little use for complex neural areas and the supraoesophageal ganglion (or brain).
The brain is a simple bilobed structure. The main sensory inputs to the supraoesophageal ganglion are the photoreceptors. The brain in pedunculates is little different
from that in other cirripedes. The ocellar nerves, which may be homologues of the balanoid antennular nerves, are separate in
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Figure 6 Pollicipes polymerus: relationship of ocellus and supraoesophageal ganglion. 1=circumoesophageal chord; 2=supra
oesophageal ganglion; 3=stalk nerve; 4=ocellar nerve; 5=photoreceptor. (After Gwilliam 1963).
Pollicipes. The function of the median nerve is not yet known in this genus but it may be that pedunculates only have the homologues of the lateral ocelli and lack the
median (Gwilliam 1987). The peduncular nerve may, therefore, be a homologue of the balanoid suprasplanchnic nerve. Gwilliam (1963), because of the activity
recorded from the peduncular nerve, suggests that it may be similar to the antennular nerve and that the ocellar nerves represent only that part of the antennular nerves
carrying lateral ocellar axons (Fig. 6).
Gwilliam (1963) examined the electrical activity of the peduncular nerves of P. polymerus. He found a rhythmical pattern of bursts of activity in light or after 15 min
in the dark. The general nature of the activity was similar although the frequency may vary. If a shadow was cast on a preparation in the light just after one of the
spontaneous bursts there was a prolonged response and then a longer than usual silence and no activity at the end of shading. Multiple shadows of 300 milliseconds
duration at 2s intervals caused a response at each shadow even after 30 such shadows. Shadows of 200 milliseconds duration and 2.5s intervals and 100
milliseconds at 1s intervals gave smaller initial responses. There was no response to shadows of shorter duration. By using shadows cast with gelatin neutral density
filters Gwilliam (1963) found that cutting out 60% of the incident light resulted in what was essentially a full shadow reflex. He also found it puzzling that the motor
output of the peduncular nerves did not adapt to multiple shadows in view of the rapid adaptation of the motor output in other cirripedes. This was not always the case,
however, for at different shadow durations and frequencies of shadow casting adaptation did occur; spontaneous rhythm may have been reset in such cases.
The rhythmic activity of the stalk nerves seen in P. polymerus but not in Lepas sp. may be a reflection of the behaviour pattern of the former species. Adults feed by
extending the cirri into a moving stream of water (see p. 340) when in water but the animals are at times exposed to the atmosphere. The regular rhythmic activity of the
stalk and the mantle motor nerves serve to provide regular pulsations of the muscles that aid blood circulation.
Photoreceptors
The structure of photoreceptors in adult cirripedes has been reviewed by Gwilliam (1987) and Anderson (1994); they do not include any work on stalked barnacles
except that
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Anderson (1994) states that pedunculates have a pair of lateral ocelli while balanids have an additional median ocellus. The photoreceptors in some pedunculates
described by Darwin (1851) and other early workers are now known as lateral photoreceptors. Pouchet & Jobert (1876), knowing that adult barnacles were sensitive
to light, and working with P. pollicipes, found that the two cerebral ganglia were scarcely distinct. From their junction arose a special nerve (the union of opthalmic and
optic nerves) and laterally the antennal nerve. The special nerve divided into three and all ended in a common mass where two pigmented spots corresponding to the
eyes were found. These two eyes were separated by a space greater than their diameter. Koehler (1889a) admits that the arrangement in balanids is more complicated
than in pedunculates. He puts Pollicipes in an intermediate position as it has an eye resulting from two organs not completely fused together. Gruvel (1893b) gives an
account of ocelli in Lepas.
There does not appear to have been any modern work on Pollicipes photoreceptors.
Endocrinology
The study of endocrinology in cirripedes has been restricted and only comparatively recently have advances been made (Clare 1987, Walker 1992).
P. polymerus has the distinction of being the first cirripede in which neurosecretory cells were demonstrated (Barnes & Gonor 1958a,b). In the central nervous
system of this pedunculate, when sectioned and appropriately stained, two types of Gomoripositive neurons were found in both the supra and suboesophageal
ganglia. The central nervous system is similar to that of other pedunculate cirripedes. It has a supraoesophageal ganglion (brain) connected by oesophageal
commissures to the large suboesophageal ganglion on the ventral nerve cord. There are four other thoracic ganglia, the last two being fused together (see p. 324). A
freshly dissected nervous system did not show the bluewhite refractile appearance seen in fresh decapod neurosecretory tissue but the suboesophageal ganglion often
looked white and opaque in contrast to the more transparent appearance of the rest of the nervous system.
Barnes & Gonor (1958a,b) found two types of neurons both containing neurosecretory granules surrounding the centre neuropile of the ganglion. The most
conspicuous cell bodies were large (30–45 μm diameter) and irregularly pyriform with abundant and strongly chromophilic cytoplasm. Their Nissl substance was
concentrated in a zone round the oval nucleus that was acidophilic and which, in fixed sections, was 12–15μm diameter. Small vacuoles were found around the
periphery of these cells which were found in all ganglia including the supraoesophageal. This has nothing to distinguish it from the ganglia of the ventral nerve. A cycle
of secretory activity was described for this larger cell type. The second type of neurosecretory cells were about 20μm in diameter and characterized by extensive
vacuolation, each cell containing a few large vacuoles that almost completely filled the cytoplasm. The contents of the vacuoles did not stain by any of the methods used.
The nucleus was spherical (6–8 μm diameter) and contained acidophilic nucleoplasm. Neurosecretory granules similar to those in the larger cells were found in the
cytoplasm and around the nucleus. No secretory activity could be distinguished in these cells.
Barnes & Gonor (1958b) were largely concerned with the central nervous system of P. polymerus but they also made sections of whole animals; these sections did
not show any evidence of storage organs. No obvious tracts containing neurosecretory material could be found leaving the chain of ganglia.
Some balanid cirripedes were also examined by Barnes & Gonor (1958a,b) and they
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found some deeplystaining granules in the major ganglia but these were smaller and less defined than in P. polymerus; this may have been related to season.
Neurosecretory cells have now been confirmed in several other cirripedes (McGregor 1967, Klepal 1985, Clare 1987 and references cited therein, Anderson 1994).
Circulatory system
The circulatory system found in thoracican cirripedes is different from the classical open system found in many crustaceans (Burnett 1987, Anderson 1994). Darwin
(1851) was one of the first to comment on the haemolymph circulation in cirripedes. He remarked that “… like others, I have failed, as yet, in discovering a heart”. He
pointed out several circulatory structures in lepadids and indicated the direction of flow through these pathways. Much of what he discovered has been confirmed in
modern work and the structures have been given uptodate names. Nussbaum (1890) gave a detailed account of his own work on the circulation in P. polymerus but
made no reference at all to Darwin’s work. He inferred similarities between a vertebrate heart and the sinuses of cirripedes. These comparisons were misleading. He
was correct in saying that the haemolymph flowed from the rostral sinus to various parts of the body and that the flow into the peduncle was through the rostral valve,
but he was wrong in assuming that haemolymph flows into the cirri from the rostral sinus. Burnett (1972) found Gruvel’s (1904a, 1905) accounts difficult to follow
because of a lack of adequate illustrations and in any case little was added to Darwin’s account. The circulatory system of the burrowing pedunculate Lithotrya was
examined in detail by Cannon (1947) and he thought that of the intertidal Pollicipes polymerus to be about the same. Burnett (1972) found that Cannon had under
estimated the complexity of the system in P. polymerus. By injecting a silicon rubber compound into P. polymerus Burnett (1972) was able to examine the system in
great detail and found that it was essentially closed. Burnett (1974, 1975) found that other barnacles had circulatory systems similar to P. polymerus but less complex.
The circulatory system in P. polymerus can be divided into four sections (Burnett 1972, 1987, Anderson 1994). The first is circulation through the peduncle and
mantle; the second is distribution of the haemolymph from the scutal sinuses to the maxillary gland, cirri and some parts of the gut. Peripheral circulation, the third part,
takes haemolymph from the cirri and scutal sinuses to the subcuticular areas of the body and other parts of the gut. The fourth section is the collective circulation by
which haemolymph, mostly from the peripheral circulations, is returned to the peduncle (Fig. 7). Burnett (1972) describes his divisions of the circulatory system in great
detail. He found that the prosomal sinus connects with the rostral vessel and that this has paired, flattened, lateral enlargements connecting dorsally with the prosomal
sinus. The ventral surface of each rostral enlargement acts as a valve (prosomal valve) by covering the passage between the rostral vessel and the prosomal sinus. The
prosomal valves can allow haemolymph to flow in either direction depending on the circumstances. Normally blood is pumped from the prosomal sinus into the rostral
vessel. If this is distended then it can flow in the opposite direction.
Anderson (1994) listed three critical aspects of the flow system from a hydrostatic skeletal point of view. There has to be the ability to inflate the capitular mantle
tissue thus separating the opercular valves. The body has to be inflated as a basis for the hydraulic extension of the thorax, cirri, and penis during respiration, feeding,
and mating, respectively. There has also to be the ability to reverse these processes and to return the blood to the peduncle.
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Figure 7 Pollicipes polymerus: diagram of direction and main features of haemolymph circulation in peduncle and mantle, as=adductor
scutorum; mk=mantle knobs; mp=mantle plexus; ms=muscle sinus; op=ovarian plexus; pv=peduncular vessel; sv=scutal valve;
sve=scutal vessel; s=sinus. (After Burnett 1987).
The location of the haemolymph pump in barnacles has been disputed for some time (Cannon 1947, Burnett 1972, Fyhn et al. 1973). In Lithotrya Cannon (1947) put
it between the adductor suctorum and the base of the oral cone, that is in the rostral sinus; he called this a “blood pump” and not a heart because the muscles were in
the sinus rather than surrounding it. According to Burnett (1972) the rostral sinus in Pollicipes polymerus did not contract in a rhythmic manner and, therefore, could
not be responsible for the continual movement of haemolymph around the body. He found that three pairs of skeletal muscles that compressed the dorsolateral channels
and prosomal sinus were responsible for pumping the haemolymph. The rostral vessel appeared to be a vestige of a “heart” in which the pump muscles had been lost.
The peduncle and mantle of P. polymerus may also be capable of pumping action. Barnes & Reese (1960) found that the peduncle was constantly contracting,
lengthening, and bending. Gwilliam (1963) recorded regular rhythmical activity of the stalk and the mantle motor nerves (p. 325). Such regular pulsations of muscles
must aid blood circulation. Burnett (1972)
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showed that contraction of the muscles of the peduncle and mantle probably pumps the haemolymph through the mantle to the scutal vessels. From there the force on
entering the scutal sinuses was enough to send it through vessels of the body. Irregular contraction of muscles in the scutal sinuses could supplement this force if
pressure from the mantle were reduced.
Besides small movements associated with haemolymph circulation the peduncle can contract when stimulated mechanically and later expand. Contraction is mediated
by the peduncular longitudinal muscles and expansion may be caused by the contraction of the circular and oblique muscles and by blood pumped into the peduncle
from the dorsolateral channels and the prosomal sinus. These channels and sinus are probably the storage place for haemolymph in the body. Contraction and
expansion of the peduncle involve rapid exchange of large amounts of haemolymph.
Cirri of adult P. polymerus do not beat as do those of many other cirripedes. During feeding the cirri remain fully extended (Barnes & Reese 1959); a ramus flexes if
a food particle stimulates it and then returns to its normal extended position. The lack of cirral beating indicates that there can be no connection with the pumping organ;
the epineural sinus is probably not connected to the dorsolateral channels. This is supported by Burnett’s (1972) observations.
The haemolymph
Very little has been done on the haemolymph of Pollicipes species. Walker (1992) mentioned haemolymph cells (haemocytes) in balanomorph cirripedes. Waite &
Walker (1988) attempted to establish the presence or absence of respiratory pigments in barnacles. They found no evidence of such pigments in the haemolymph of
balanomorph barnacles. This was in agreement with Petersen et al. (1974) who could find none in Pollicipes polymerus and Innes (1985) who could find none in
Calantica spinosa (=Pollicipes spinosus). The effects of aerial respiration and acclimation temperature on the in vivo acidbase status of haemolymph in Calantica
was examined by Innes (1986). He obtained a similar result to that found by Petersen et al. (1974) in Pollicipes polymerus. An increase in haemolymph of the
haemolymph exceeded 5mmHg. When submerged in wellaerated sea water at “normal” acidbase status the peduncular haemolymph maintained a constant relative
alkalinity of 0.2pH units (Petersen et al. 1974). This may be connected with the lack of a respiratory pigment in the haemolymph. In Capitulum mitella (=Pollicipes
mitella), the body fluid was found to contain a nonspecific haemagglutinin (Kamiya & Ogata 1983).
Waite & Walker (1992) determined total lipid and protein levels in the haemolymph of a balanid and compared them with those in a pedunculate, Lepas anatifera.
The neutral lipid and phospholipid were about the same in each, but lipid was much lower in the pedunculate than in the balanid. Florkin & Blum (1934) obtained blood
by puncturing the base of the peduncle of Pollicipes pollicipes and found it to have a protein content of 0.2%; they gave a typical average value for crustaceans as 4%.
The carotenoids and carotenoproteins present in both the body and peduncular fluid of adult P. polymerus and of various stages of the egg lamellae have been
investigated by Holter
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Table 4 P. polymerus: relative percentages of the pigments isolated from mature and immature bodies, peduncular fluid, and three stages of egg
lamellae. (After Holter 1969).
Pigment, %
Material tested ß carotene Isozeaxanthin (?) Astaxanthin ester Lutein Zeaxanthin Astaxanthin
Mature body 4 2 26 34 5 29
Immature body 6 2 25 32 4 31
Peduncular fluid 3 1 3 33 3 57
Egg lamellae, early 8 – 28 36 5 23
Egg lamellae, medium* 11 – 20 32 2 33
Egg lamellae, late* 10 – 8 20 3 60
* These percentages add up to 98 and 102, respectively, but are as given by Holter (1969)
(1969). The egg lamellae were staged according to colour and compared with the stages of development given by Barnes (1965). The carotenoids were extracted with
acetone and initially separated chromatographically. The results were confirmed by comparison with known carotenoids. Protein complexes were extracted with 0.2M
phosphate buffer, pH7, precipitated with ammonium sulphate and eluted on columns of DEAEcellulose. Six carotenoid pigments were found in the bodies and
peduncular fluid but only five in the egg lamellae; isozeaxanthin(?) was not found in the egg lamellae (Table 4). There was no difference in the kind or relative amount of
carotenoids in adult mature and immature bodies, suggesting that P. polymerus may not metabolize its large amount of astaxanthin from carotene percusors as no
intermediates were found. This does not mean that in this barnacle metabolism may not take place rapidly in the larval stages. The astaxanthin may, however, be taken
directly from food sources. In the peduncular fluid and the egg lamellae the astaxanthin was bound to a highly unstable protein (probably a lipoprotein).
The characteristics of the peduncular fluid, and particularly its contained protein complex, were unusual. The same carotenoids as those in the body were found but
the astaxanthin was bound in some way to what could be a lipoprotein. Holter (1969) suggested that because developing egg lamellae contained a similar astaxanthin
lipoprotein the peduncular fluid might be used in some other way during embryo development. This seems doubtful. The more likely source is the lipoprotein complex
or the esterified astaxanthin of the early stage embryos.
Ninhydrinpositive substances (NPS) of the muscle tissue of P. polymerus have been determined by Fyhn et al. (1972) at different haemolymph osmolalities. They
found that the variation in NPS concentration was probably a passive result of variation in the volume of the muscle cells. At an osmotic coefficient of one, NPS could
account for approximately 20–25% of the intracellular osmolality. Muscle tissue from animals exposed to air at 26°C and 55% relative humidity for varying times (up to
15h) was also examined. It appeared that exposure of about 8h was needed to cause any increase in NPS concentration. The osmolality and NPS concentration of the
haemolymph and the relative water content and NPS concentration of the muscle tissue were determined in the natural environment. The animals were tested at
different times during a tidal cycle; they were all of a similar size and from the same cluster at a +5ft tidal height. The weather conditions during the experiment varied
from sun to overcast, light to heavy rain, and wind to no wind. The air temperatures were 10–19°C and the seawater temperature was 9°C. The osmolality of the
haemolymph fluctuated according to the weather conditions during exposure to air. The relative water content of the muscle after air exposure of 9h showed an increase
of about 2% compared with the value for submerged animals. It was a wet day, with particularly no wind, and an air temperature of 10–11°C.
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Otherwise there were no significant variations throughout a tidal cycle.

In P. polymerus the volume changes of the muscle cells were less than expected but the variation in NPS concentration in the muscle tissue could be accounted for
by the change in cellular hydration. Intracellular solutes other than free amino acids must, therefore, be involved in regulating intracellular osmolality. The field results
support this. A reverse relationship of NPS concentration of the two compartments (which was not found) would result if there were any interchange of amino acids
between muscle cells and haemolymph. Values for the muscle tissue of exposed animals may have been affected by the time needed to change the intracellular amino
acid concentration. The time needed to show an increase in NPS concentration was more than 8h and air exposure of P. polymerus in a natural population lasts for a
maximum of 9h. This is added evidence that amino acids can be excluded from the regulation of intracellular osmolality during intertidal periods.
Osmoticallyactive particles can be removed from the body with the fluid. Fyhn et al. (1972) found that mechanical stimulation of animals exposed to air caused fluid
to be expelled through the aperture of the capitulum after a delay of 1–2min. Samples of the fluid were collected from animals after 4–5h exposure to air. Haemolymph
samples were taken from the same animals after collection of the excreted fluid. The osmolality of the excreted fluid was virtually unchanged at least up to 75min after
the disturbance. It was 20–25mosmoles hyperosmotic to the haemolymph of the animals but this may have been because there was some evaporation of the excreted
fluid before sampling. It is probably better, therefore, to assume that the excreted fluid is isosmotic to the haemolymph. This means that the fluid can have no
osmoregulatory function. A possible use may be to keep the respiratory surface wet during aerial respiration (Petersen et al. 1974) and to keep egg lamellae bathed by
a suitable medium during exposure to air. The flow of excreted fluid increases as the animal’s peduncle contracts and so expulsion may be the result of high hydrostatic
pressure (Fyhn et al. 1973).
Haemolymph pressures
A high haemolymph pressure such as is produced in a closed haemocoelic system is necessary in an animal such as P. polymerus in which a turgid peduncle has to
support a capitulum in an elevated position. This was emphasized by Burnett (1987) and Anderson (1994). Fyhn et al. (1973) made continuous readings of the
haemolymph pressure in whole and unrestrained P. polymerus by inserting catheters into the body sinuses. The pressures were recorded with Statham pressure
transducers and a Beckman oscillograph. They found pressures ranging from 110 to 350cm H2O above atmospheric pressure in the peduncle as well as in the
capitulum. The pressures were pulsatory and rhythmical although the pulse amplitude and frequency varied considerably. The pulse pressure and absolute pressure
(mean of systolic and diastolic pressures) were stable for up to 3.5h (time of experiment) when an animal was not distributed. When mechanically stimulated the
peduncle immediately contracted and the pressure increased to 500cm H2O. Pressures were independent of the weight of the animal between 3 and 35 g. Presumably
this was the weight of peduncle and capitulum; the authors did not specify. Changes in body temperature did, however, affect the pressure. At 18–22°C the absolute
pressure was only about 70% of that at 7–17°C.
The amplitude of pulsations was higher in the capitulum (15–70 cm H2O) than in the peduncle (usually 8–10cm H2O but ranging from 3–20cm H2O). The
contraction frequency increased with body temperature; it had a Q10 of 5 to 10 between 7 and 15°C and a Q10 of 2 to 3 between 15 and 25°C. The pulse amplitude
was independent of temperature. The con
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Figure 8 Pollicipes polymerus: duration of systole (A) and diastole (B) of contractile pumping mechanism as a function of contraction
frequency. (After Fyhn et al. 1973).
traction phase (systole) and relaxation phase (diastole) of the pressure pulses at different contraction frequencies are shown in Figure 8. The greatest dependency on
frequencies was at <10 beats min−1 and systole duration varied much less with contraction rate than did diastole duration. Systole made up an almost constant
proportion of the cycle length (20–27%) at different frequencies between 10–50 beats min−1.
The rhythmical haemolymph pressures recorded in P. polymerus were much higher than those usually found in invertebrates. The pulsatory pressure suggests
unidirectional muscular pumping of the haemolymph in a system of high peripheral resistance.
Waite & Walker (1986) also measured haemolymph pressure in P. polymerus by inserting a catheter directly into the peduncle. The absolute pressure recorded,
admittedly on a single animal, was 74.3cm H2O at 14.8°C, much lower than reported by Fyhn et al. (1973). The pulse pressure was, however, 11.5cm H2O which
does compare with Fyhn et al.’s figure of 8–10cm H2O. In a comparative study of haemolymph pressures in cirripedes Walker & Anderson (1990) quote average
figures for three P. polymerus credited to M.E.Waite (unpubl.) as 38–300cm H2O for the absolute pressure, 5–15cm H2O for pulse pressure with 6–11 pulse
frequencies per minute at 8–9°C; a provoked pressure was given as 240–430cm H2O. Unpublished results of G.Walker for three P. pollicipes are also given in
Walker &
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Anderson (1990). These are 155–245cm H2O for the absolute pressure, 6–10cm H2O for pulse pressure with 3–9 pulse frequencies per minute at 8°C. A provoked
maximum pressure was given as 200–246 cm H2O and a natural valve closure as 197–246 cm H2O. Waite & Walker (1986) reconciled the high haemolymph
pressure of P. polymerus with the force required to extend the peduncle. Unfortunately, when Anderson (1994) was discussing haemolymph pressure and quoting the
already published results mentioned above, he gave the units as mm H2O instead of cm H2O.
Strength of peduncle
Crenshaw (1979) considered the peduncle of P. polymerus to be a pressurecontaining cylinder that had to support the capitulum. The function of the pressure of
haemolymph in the peduncle was to provide hydrostatic support for the organism. This hydrostatic pressure would produce tension in the walls of the peduncle and if
they were to yield the support system would collapse. Crenshaw made mechanical tests on the integument of the peduncle by stretching pieces immersed in sea water.
He used pieces cut longitudinally and circumferentially from living barnacles. The integument continued to stretch for as long as the tests were run (3.5h). Compliance of
the integument was the same in both the longitudinal and circumferential directions. The compliance of a cylinder of integument when tested in the longitudinal direction
was higher than in either set of flat test pieces. Cylindrical pieces were, therefore, assumed to be better estimates of the compliance of the integument. Pressure in the
peduncular sinus was not correlated with the stretching of the integument. Hydrostatic pressures were maintained when the peduncle was very stretched as well as
when contracted. Thick sections of the integument examined with a polarized light microscope showed that it consisted of birefringement laminae. Tests showed that the
integument contained chitin. Networks of birefringement anastomosing connective tissue fibres were found throughout the whole peduncle. Tests proved that these
contained protein. The fibre network ran between the muscles, sinuses and ovary in the peduncle (see also Burnett 1972). The fibres branched frequently and yet were
continuous. In transverse section they appeared as spokes going radially from the centre to the inner wall of the integument, which they penetrated, and terminated in its
inner laminae. Garrault (1934) and Mahmoud (1959a) found a similar arrangement in P. pollicipes. Koehler (1889a) found the connective tissue of P. pollicipes
particularly dense giving the peduncle added resistance. According to standard engineering practice, Crenshaw (1979) did not believe that a cylinder of dimensions
similar to the peduncle would be capable of acting as a pressurecontaining vessel; the cylinder would deform into a sphere. Such deformation was not found in live
barnacles at high haemolymph pressure. In the peduncle of P. polymerus there are three muscle layers involved in containing the pressure, longitudinal, oblique, and
circular. When contracted these muscles apply a force against hydrostaticallygenerated tension in the integument and so relieve the stress. The anastomosing
connective tissue fibres may also reinforce the integument. Fibrous protein such as collagen and elastin can withstand tension.
The structural arrangement of the pressure container in a cirripede peduncle may be different from that in most hydrostaticallysupported organisms but the functional
role is similar. Burnett (1987) suggests that the muscles of the body and peduncle act together to produce haemolymph pressure high enough to give rigidity to the
peduncle while at the same time allowing it to bend.
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Excretion
The excretory organs of adult barnacles are paired maxillary glands situated in the anterior part of the body, below and to the sides of the foregut, each with an opening
at the base of the maxilla. The organs include three parts: the end sac (mesodermal), the efferent duct (endodermal), and the terminal duct (exodermal) (White 1987,
Walker 1992, Anderson 1994). In pedunculate barnacles Darwin (1851) first described paired sacs lying in the anterior part of the body but he did not recognize them
as excretory organs. Hoek (1883) was the first to realize their excretory nature. Other workers followed and excretory organs in many different adult barnacles have
now been described morphologically (White 1987). These include Lepas (Gruvel 1894). P. polymerus (Nussbaum 1890) and P. pollicipes (Koehler 1892, Gruvel
1893a,c, 1904a, Bruntz 1902, 1904). Later work on additional species, but not on Pollicipes, was reviewed by NilssonCantell (1921). Batham (1945) described the
organs in Calantica spinosa (=Pollicipes spinosus). Modern histochemical and ultrastructural techniques have been applied to some balanids (White & Walker 1981)
and to Ibla (Klepal 1985).
The relative proportions of the three parts of the system vary with species. In Pollicipes the end sac is large and the labyrinth much reduced. Koehler (1892) thought
the end sac in P. pollicipes was divided into a number of compartments but according to Bruntz (1904) this divided appearance is due to the extremely loose character
of the epithelium. In balanids White & Walker (1981) found the excretory organ to be surrounded by a network of elastic fibres and suspended within the body by
connective tissue and muscle connections. The end sac is supplied with haemolymph from the scutal sinus; this agrees with Burnett (1972) for P. polymerus. The
supply of haemolymph to the efferent duct is more restricted although it improves towards the terminal duct.
In most crustaceans an important function of the excretory organs is ion and haemolymph volume regulation. Another is the removal of nitrogencontaining
compounds arising from catabolism of proteins and nucleic acids. It is generally accepted that the nitrogenous endproducts are excreted largely as ammonia, although
considerable aminonitrogen, some urea and uric acid may be formed (White 1987). The efferent duct has been found to concentrate copper when animals live in
environments with high levels of trace elements. The copper is normally associated with sulphur. The parenchyma surrounding the excretory organ has also been shown
to accumulate trace metals (White 1978). This ability led P. polymerus to be one of the animals used by Shokes & Callahan (1978) in their study of the potential and
adverse effects of drilling activities and anthropogenicity on marine organisms. Mix et al. (1979) made preliminary studies to evaluate the potential of using the larval
stages of P. polymerus in marine bioassays.
Orientation and movement of peduncle

P. polymerus does not usually extend into the upper parts of the intertidal zone although there are a few exceptions. It normally occurs below Chthamalus species and
the main Balanus glandula belt. Observations made by diving indicates that it does not occur sublittorally. It is not usually found on the lower parts of pier piles, or on
sublittoral rocks or reefs even in areas where it is abundant intertidally. It has already been mentioned that the animal favours growth in groups (p. 305 and Fig. 1). The
form of the aggregate depends largely on both the degree of exposure and the microenvironment. Cyprids are attracted to these groups and
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juveniles prosper there as they gradually extend their peduncles to attach on to the primary substratum (see p. 356). Examinations of these groups of animals show that
over a restricted area most of the individuals have their capitula orientated in the same direction (Fig. 9; Barnes & Reese 1960). Except in places where wave action is
relatively light it is evident that, in contrast to the reactions of the cirri of operculates to currents, the dominant orientation is not to bring the anterior face of the cirral net
against incoming waves. On the contrary, it is usually such as to bring the open cirral net at right angles to the flow of water but with the anterior or captorial face into
the stream as it pours off the rocks subsequent to the breaking waves (see p. 337). The shape of the capitulum, once this orientation is determined, allows the animals
to form packed masses, whose dominant orientation is readily seen. Such orientation is not to the shoreline or the general wave direction but is determined by the
microtopography of each local situation, varying with any irregularities of the substratum and its position with respect to the breaking waves. The anterior face of each
cirral net is brought to face the rush of water, in the one case as it surges down the seaward side of a boulder after the wave has broken, in the other (at a later stage of
the tide) as it flows over and pours down the landward side. Where there are well marked channels either in the rocks themselves or formed by a series of contiguous
boulders, the dominant orientation is to the axis of the channel with the cirral net at right angles to it and the captorial face directed to the current as it flows down the
channel after a wave has broken. Animals on the sides, as distinct from the floor, of such a channel, are similarly orientated but the capitulum as a whole retains the
same relative position to the rock surface. Where, as a result of microtopography, there is a general turbulent motion rather than a distinct pattern of flow, then there is
no dominant orientation; such is often the situation at the corners of large boulders or amongst a collection of small rocks.
It has been shown that P. polymerus has no regular cirral beat (see p. 340) except in juveniles (see p. 342). In its macrofeeding activity, food is carried to the
mouth after the appropriate stimulation of the individual cirri (Barnes & Reese 1959). In the absence of a rhythmical beat the orientation with respect to water currents
is an adaptation that ensures a favourable position for feeding activity. When the anterior face of the cirral net is at right angles to the dominant water flow then both the
maximum chance of contact with planktonic material and the greatest efficiency in its transport by the cirri to the mouthparts are ensured. It is not at first clear, however,
why this leads to the anterior face of the cirral net being directed to the backwash of water rather than to the incoming waves. In P. polymerus a certain amount of
general water movement is essential before more specialized responses are readily evoked. Under natural conditions it would seem that this primary level of activity is
initiated and then in part maintained by the incoming waves, so that the animal is then in a more favourable state to respond to directional wave currents.
P. polymerus may be maintained for quite long periods in the laboratory, healthy, and actively feeding; this is best done by playing a continuous jet of water over the
animals while maintaining them under running water, rather than by stirring or bubbling with air. When such a jet is directed laterally on to the cirral net of an active
animal it slowly turns so that the anterior face of the cirri is brought towards the jet. This orientation is not brought about by a temporary twisting of the cirri alone but by
a slow turning of the whole capitulum. When the jet is directed at the peduncle, no turning takes place, although with a strong jet the peduncle may bend towards it so
that although the receptors to unidirectional water movements appear to be situated in the cirri there must be integration with the ultimate response in the musculature of
the peduncle. If the valves are tied together with a piece of twine, no turning response is obtained to a jet of water but it is evident that there is an effort to open the
valves as the
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Figure 9 Pollicipes polymerus: small areas showing orientation to water flow. Arrows indicate direction of backwash of waves.
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Figure 10 Pollicipes polymerus: diagram showing change in orientation of individuals in a group caused by a water flow (W) of 1 litre
per six seconds from two jets of 5mm bore and 12mm apart placed 12cm from the leading edge of the group of animals. Note
change in orientation of animals unless the opercular valves were prevented from opening by tying them together (in black).
(After Barnes & Reese 1960).
twine may be considerably stretched (Fig. 10). Receptors responding to general water movement rather than directional flow may, therefore, be present in the
peduncles. When the twine is removed, the capitulum opens, the cirri are extruded, and the turning response takes place. With an isolated individual this turning
response is easily obtained; under the stimulus of even a moderate jet directed laterally on to the side of an animal whose cirri are expanded, it slowly turns through 90°.
Although the evidence strongly suggests that the effective receptors that initiate the turning response are in the cirri, it is difficult to be quite certain that this is the case.
Under a vigorous flow of water and with a fullyactive animal the body is freely exposed to water, being raised in the mantle cavity. It is possible, therefore, that the
receptors lie within the body surface. Experiments in which a very fine jet was directed at the cirri tended, however, to negate this suggestion because under these
conditions the body is largely protected from the impact of the water by the valves and yet the response is still obtained. Attempts to resolve the problem by observing
animals in which one or more cirri had been removed were inconclusive. There are nerve endings at the base of the spinous processes of the cirri and these may be
directly involved in the response. Whether these receptors are identical with those responsible for the “feeding” responses of the cirri to touch is not known but because
a different reaction is obtained, it seems possible that a different set of receptors is involved.
In P. polymerus the reception of the turning stimulus by the cirri is believed to be followed by integrative action, orientation being achieved by rotation of the whole
capitulum by means of the muscles in the peduncle. The rotation is, therefore, essentially “temporary” and whereas it may be maintained virtually constant where
unidirectional currents predominate it does not become fixed by shell growth as in an operculate. This would seem to have considerable advantages in situations where
the water flow, although dominantly unidirectional, is subjected to small and variable changes resulting from variations in the angle of approach of the waves;
adjustments of the cirri can readily be made to such changes.
When a group of animals still attached to their substratum is observed in still water or nondirectional currents, either in the field or the laboratory, they show little
sign of activity but from time to time the valves open and the cirri are partially extruded. There is no macrofeeding activity and under these conditions the cirri are rarely
expanded sufficiently to feed. This has been effectively demonstrated by Barnes & Reese (1960) using kymograph records and timelapse photography. Lewis (1981)
quotes unpublished work of S.Donaldson & D.J. Prior recording a nervous activity at the end of a severed cirrus in an adult P. polymerus. No
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such activity was recorded from juveniles. The spontaneous impulses in adults may represent a remnant of the cirral beating pattern of juveniles as the adult nervous
activity was about the same as the rate of cirral beat in juveniles of 1–2 mm RC. It does not seem possible that this rhythm represents the behavioural rhythm observed
in adults by Barnes & Reese (1960) because the movements only happened every 50–100s. If the water is stirred vigorously, the activity of the animals is increased;
the valves open more widely and the cirri may be fully expanded. The behaviour is similar on the shores; as the tide rises the water reaches the animals, the cirri are first
protruded and hang out somewhat limply between successive waves. With increasing exposure to wave action, the cirri become fully expanded and activity of the
peduncle in the form of bending movements often takes place. It is very evident that for effective cirral response to various forms of stimulation a considerable amount
of water movement must be present to maintain a “basal” activity.
Not a great deal is known about the behaviour of the cyprids of P. polymerus as regards their distribution at settlement and any interpretation must, therefore, be
tentative. By analogy with other intertidal cirripedes it might be expected that they would tend to remain in the upper water layers and that settlement would be greatly
influenced by wave action. There is little doubt that settlement is heavy among adults of the same species. Whether species recognition is involved in this behaviour or
whether it is only the specialized requirements of the surface texture that are met most commonly by the surface of the peduncles of adults, is not known. In any event
the phenomenon is clearly responsible for the maintenance of dense masses of the animal in suitable places with the advantage detailed on p. 358.
The restriction of P. polymerus to places with wave action is clearly related not only to the necessity for a certain general stimulation to initiate full activity, but also to
the method of macrofeeding, once this level of activity has been reached (Barnes & Reese 1959). The ability to respond to unidirectional currents by a twisting of the
peduncle allows full advantage to be taken of local macro and microtopography in feeding activities. The fact that the animals are orientated to the runoff waves over
and between boulders, rather than the initial breaking wave, permits a longer feeding time. In quiet places such as protected bays or harbours as well as in the sublittoral
the necessary stimulation is absent and Pollicipes do not occur.
There is less information regarding the factors limiting the species at the upper level of the intertidal zone. Two important factors may be suggested. First, because
rather specialized conditions are required for full feeding activity, and because the conditions even in the most favourable sites are only completely satisfied during a
limited part of the tidal cycle (optimal only from the time the tide reaches the animals to when they are completely covered) this period, at the upper levels of the shore
may be insufficient to sustain adequate growth. Secondly, desiccation at the upper tidal levels may be too severe for both spat and adults. As in intertidal balanids,
water loss from the body is in part controlled by adjustment of the capitulum, but in a pedunculate water may also be lost from the peduncle, and over such losses the
animal has only a limited control, achieved by a contraction of the peduncle to reduce the area of the surface exposed (see p. 348). It seems unlikely that competition
limits the upward spread; this could have come from Balanus glandula and it is difficult to see how this species, in view of the “adjustments” of the peduncle in
Pollicipes polymerus, could successfully prevent the upward penetration of the latter species.
Over the greater part of the west coast of North America the distribution of P. polymerus within the intertidal is that already described. There is one notable
exception which provides an interesting test of the reliability of the ideas concerning behaviour and distribution. The waters of the San Juan Archipelago are relatively
protected and, compared with the outer
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Pacific coast, wave action is very slight. P. polymerus might be expected to be absent but, as has been seen (p. 306), this is not the case. The species is by no means
as abundant as on the outer coast and indeed rarely forms dense masses on vertical faces of rock. On parts of the coast, however, as a result of the nature and
configuration of the rocks, the steep cliffs are often cut by gullies that extend well above hightide level. It is in such gullies that the pedunculate is common, and here it
may form quite dense masses that are particularly clearly orientated to the direction of water flow down the gullies. With moderate or even light wave action
observations show that the breaking waves are channelled with some force up such gullies to give a directional return flow of water. Turbulent conditions and water
flow are, therefore, adequate for Pollicipes. Furthermore, with the water channelled in this way, a band of animals within the gullies often extends over the whole
intertidal range so that at the upper end of the gullies Pollicipes is present across the levels occupied by Balanus glandula and Chthamalus dalli and even at higher
levels than these species reach on the open rock faces.
In Europe Pollicipes pollicipes is also associated with exposed situations and turbulent water conditions. Several workers record the cirripede being found in deep
channels in steep cliffs characteristic of the exposed west coast of Brittany (see p. 307). It has also been found in caves when waves break with extreme violence at the
entrances and the water is forced into the caves and rushes out again as seen in steep channels on San Juan Island where P. polymerus prospers (see above). Although
not examined in detail the feeding of adult P. pollicipes follows the same sequence of events as seen in P. polymerus after a change from feeding by cirral beating in
the juveniles (Hui 1983b). As the ecology of the two species is comparable the similarity in feeding behaviour is to be expected.
Williams (1991) observed bending and twisting of the peduncle of P. pollicipes and found it virtually similar to that in P. polymerus. There were some differences.
In undisturbed groups of animals there was little or no response to shading. When mechanically stimulated, however, cirri were immediately withdrawn and the capitular
valves closed, with a slight contraction (2–3mm) of the peduncle (> 45mm length). Peduncles of this length often show slight bending movements. The degree of cirral
retraction also varied. If fully extended cirri are mechanically stimulated there is partial retraction of the cirri, but reemergence occurs slowly after 5–10s. In animals
with only onethird to twothirds of the cirri extruded there is no reaction to stimulus.
Cirral activity and feeding

The principal activity of sessile barnacles involves the six pairs of cirri and the associated movement of the shell plates and mouthparts. In pedunculates there is some
movement of the peduncle (Barnes & Reese 1960). Cirral activity is necessary for ventilation of the mantle cavity and the collection of food particles. It also allows very
small particles to be filtered from the water as the current passes through the mantle cavity (Anderson & Southward 1987, Anderson 1994). Darwin (1851, 1854)
described the cirral activity and the coordinated movement of the body, but he was not correct in some of his assumptions about the musculature of the cirri and the
mechanism by which they furled and unfurled. Later workers have elucidated this. Nussbaum (1890) examined P. polymerus but Cannon (1947), working with the
pedunculate Lithotrya, did not always agree with him. Cannon (1947) described only flexor muscles in the cirri of Lithotrya and Pollicipes but no extensor muscles.
He suggested
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that extension of the cirri during feeding was due to hydrostatic pressure (see also Burnett 1987, Gwilliam 1987).
The adult P. polymerus does not have a normal cirral beat in the generally accepted way. It uses the currents caused by wave action for feeding (Barnes & Reese
1959) and keeps its cirri fully extended and rigid during feeding (Burnett 1972). After flexion caused by mechanical stimulation or a food particle the flexed ramus
immediately returns to its normal extended position. There is no detectable movement until the next stimulation. As there is no rhythmic pumping action the epineural
sinus is probably not in communication with the dorsolateral channels.
When the adult P. polymerus has extended cirri with the valves of the capitulum open harsh stimulation, such as a predator (Barnes & Reese 1959), causes the
body to be rapidly pulled into the mantle cavity, the capitular valves close, and the peduncle immediately shortens to between 10–20% of its original length. At the same
time haemolymph from the peduncle enters the mantle circulation system in larger quantities than usual. This excess haemolymph enters the scutal sinuses and reaches
the rostral sinus. The haemolymph probably enters the prosomal sinus and dorsolateral channels through the prosomal valves and it is stored there until the next
elongation of the peduncle (Fig. 11). Burnett (1972) suggested that if the rostral sinus did not act as a place to absorb the sudden influx of haemolymph caused by such
drastic movements of the animal, the adductor scutorum would not be able to contract as rapidly as it does.
Foster (1978) has given a general description of cirri and their spines and this applies to many cirripedes. Cirri and spines of P. polymerus have been described by
NilssonCantell
Figure 11 Pollicipes polymerus: diagram showing probable route of haemolymph during contraction (A) and elongation (B) of the
peduncle. In A the prosomal valves open due to high pressure in the rostral vessel. In B somatic muscles force excess
haemolymph out of the prosomal sinus; the valve in the oral cone prevents a return to the body via the rostral sinus so it returns
to the peduncle via the peduncular vessel, as=adductor scutorum; c=cirri; es=epineural sinus; me=mantle circulation; mp to
ss=mantle plexus to scutal sinus; oc=oral cone; p=peduncle; pev= peduncular vessel; ps=prosomal sinus; pv=prosomal valve;
rs=rostral sinus; rv =rostral valve; rve=rostral vessel; v=valve. (After Burnett 1987).
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(1921). A careful description of the cirral action during feeding in P. polymerus has been given by Barnes & Reese (1959). The details include the positioning of the
spines and hairs on different segments of the cirri and their pedicels; it is impossible to enlarge or add to their descriptions. No quantitative observations were available
but from experience these authors believed that the anterolateral face of the cirrus, well supplied with long spines, was more sensitive than the posterior face with
relatively short spines. It also seemed that there was a directional effect because stimulation was more effective if the spines were bent in a proximal direction. If it is
assumed that the spines are pressure receptors then there may be some structural asymmetry in relation to sensory neurons. Repeated stimulation was found to cause
fatigue.
The activity of P. polymerus is related to water movement (Barnes & Reese 1960). Extension of the cirri and their reaction to stimuli depend on water flow above a
certain critical level. Speed as well as turbulence of the water and the effect of mechanical shock such as in the surf zone all play a part. Depending upon the intensity
and subsequent events a whole series of reactions take place. When a food item strikes the distal end of cirrus VI it is rapidly brought down to between the basal pads
of the anterior cirri. If the prey tends to struggle and tries to escape more of the cirri may curl towards the mouth region to assist in preventing such an escape. When
they come together and their spines overlap, the net acts as an extremely effective captorial device. Once trapped between the bases of the anterior cirri rapid
reciprocal movement of these parts is initiated. Such “reciprocating” movements are also seen in the posterior rami of cirrus I with a relatively short toandfro
movement giving the appearance of reciprocal movement in one plane parallel to the top of the oral cone. Material seen to be extruded between these first cirri is being
rejected and must be carried over the mouth parts. These remain inactive during the rejection of material.
It is difficult to obtain direct evidence of how food selection is made. It may involve chemoreceptors that in the presence of unwanted material inactivate the
mouthparts. Rejection of material was noted by Gruvel (1904a, 1905) and Thomas (1944). Gruvel found that the labrum and outer maxillae had unicellular glands and
he suggested that food wiped from the cirri was formed into a “bol alimentaire” that was then swallowed. Thomas (1944) found glands near the bases of the first three
pairs of cirri but thought they were too far away from the mouth to function as Gruvel (1904a, 1905) suggested. Thomas (1944) called them suboesophageal
tegumental glands (see p. 320) and thought their secretions were used to collect undesirable particles and to form them into a rejection mass. It is difficult to decide
which of these hypotheses is correct. Barnes & Reese (1959) have shown that large particles are wiped off the cirri of P. polymerus and are carried forward to the
mouth; if the particle is to be rejected then it may be carried further forward over the mouthparts and rejected. So far there seems to be no knowledge of the stimulus
deciding acceptance or rejection; it is difficult to avoid the implication of a chemotactic response.
The transference of food to the oesophagus has been described by Barnes & Reese (1959). Particles are carried from the bristles of the pedicels of the anterior cirri
by the hammerlike movements of the outer maxillae. Particles are then taken from these by the inward and downward movements of the inner maxillae and mandibles
and sent towards the oesophagus (Fig. 12). A reversal movement of the inner maxillae and mandibles causes the material to be vigorously ejected from the oral cavity.
It has been stressed that the captorial feeding method described above applied to adult P. polymerus; they depend on exogenous mechanical stimulation, such as
waves or moving prey, for feeding. Juveniles respond to the same stimuli but also beat their cirri without any external stimulation (Lewis 1981). The cirral beating in
tanks of still water was similar to,
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Figure 12 Pollicipes poly merus: schematic diagram of mouthparts and their movements; the outer maxilla moves on a midline and the
inner maxilla and mandible move downwards and inwards in a curve following the wall of the supraoesophageal cavity (SOC).
P=palpus; OMx=outer maxilla; Mx=maxilla; Mn =mandible; 1, 2, 3=order of movement. (After Barnes & Reese 1959).
but slower than that of balanids; the extension took about 80% of the time required for a complete beat compared with lower values in many other cirripedes
(Anderson & Southward 1987). In fast currents the juveniles extended their cirri as in adult animals. The number of juveniles actively beating depended on
environmental conditions. In calm water in laboratory conditions 44% were beating immediately after collection, 34% after 24hours in sea water, and 27% when
immersed after exposure to air for 24hours. The beating rate dropped from 1.4–2.7 beats min−1 when food was present to 1.2–1.4 beats min−1 when starved. If cirral
beating were merely a respiratory function more animals out of their normal exposed habitat would have been expected to beat. The frequency of cirral beating was
temperature dependent increasing from about 3 beats min−1 at 10°C to about 10 beats min−1 at 15°C. At constant temperature the beating rate was inversely
proportional to the size of the barnacle decreasing from about 4 beats min−1 for an animal of 1mm RC to 1 beat min−1 at 10mm RC. Between 10 and 12mm RC
beating was only intermittent and at ≥14mm RC there was none.
A similar behaviour has been demonstrated in P. pollicipes from the west coast of France by Hui (1983b). Three juveniles all 3mm RC showed cirral beating
activity. The cirri emerged within 3–6min of immersion and beating started 2min later. The rate of beating ranged from 2.3–5.5 beats min−1. The extension phase took
on average 51% of the time required to complete a beat compared with 80% in P. polymerus (Lewis 1981). The cirral activity also differed somewhat. The cirri in P.
pollicipes were not extended fully but were simply unfurled and moved in and out of the aperture of the capitulum. Only a few juveniles were available to Hui (1983b);
a more extensive investigation might yield slightly different results. The comparable behaviour of the two species is, however, obvious.
Mouthparts and digestive system

The mouthparts for the Pollicipes species are described in Table 2 (p. 315). The anterior mouth field is guarded by the labrum, which protrudes in may pedunculates,
as it does in
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Figure 13 Diagrams of mouthparts of Pollicipes pollicipes (A) and P. polymerus (B). a=mandible; b=maxilla; not to scale. (A, after
Broch 1927; B, after Cornwall 1925, 1955).
Pollicipes, and is termed “bullate”. The crest of the labrum bears small teeth. On either side of the labrum are the mandibular palps guarding the anterior ventral mouth
field. The pair of mandibles are toothed and are directed towards each other. The pair of maxillules (1st maxillae) are also bladelike and lie parallel to, and behind, the
mandibles. The maxillae (2nd maxillae) are triangular lobes forming the posterior margin of the mouth field. The maxillary gland opens at the base of the maxilla. On
either side of the oral cone are the first pair of thoracic limbs (cirri I) modified as maxillipeds. These are shorter and more densely setose than the remaining cirri and
function in food transfer from the other cirri to the oral cone (Petriconi 1969, Foster 1978, Anderson 1994). Nussbaum (1890) made a detailed study of the
mouthparts of P. polymerus. Broch (1927) described those of P. pollicipes and said that they were similar to those of P. polymerus (Fig. 13). Krüger (1911b) and
NilssonCantell (1921) have also described mouthparts.
The digestive apparatus socalled was known to early workers (Gruvel 1893a, 1904a, 1905) and Nussbaum (1890) describes it and the associated glands in P.
polymerus. He also gives dimensions of the main parts of the alimentary canal and cell and nucleus diameter of some of the gland cells. Some of these dimensions do,
of course, vary with the size of animal. The most recent work on the alimentary canal or tract of a pedunculate cirripede must be that of Klepal (1985) on Ibla. She
describes the various parts—mouth and foregut, midgut, and hindgut—including the musculature. The muscle system of Ibla and other pedunculates are generally
similar (Anderson 1994).
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Stomach contents
The stomach contents of barnacles reflect to some extent the environment from which they came, their mode of feeding, and the food available. Often there is a lot of
debris present and particulate matter <2μm diameter may be taken. Such small particles cannot, however, be taken by the captorial activity of the cirri. Fine material
can be filtered from the water in the mantle cavity current by the fine spines and hairs at the base of the cirri. Secretions of the glands there help to bind the particles
before they are carried forward to the mouthparts to be accepted or rejected.
Captorial feeding is the characteristic method in adult P. polymerus as seen above (p. 340). In most stomachs of P. polymerus there were only a few sand grains,
very few diatoms, and only a small amount of fine particulate debris. Most of the contents were crustcean remains 500–1000μm long. Small copepods were common
as well as cirripede nauplii and cyprids (Barnes 1959).
Howard & Scott (1958) demonstrated that P. polymerus could capture and ingest prey originally longer than the digestive tract of the barnacle. The prey, trapped
by an individual cirrus was passed towards the mouth where the prey was “bitten in pieces, folded or compacted to fit the gut”. These authors made regular
examinations of the gut contents for a year and found the usual diet of P. polymerus to consist nearly always of copepods, algae and unidentified particulate matter,
cirripede moults, cyprids, amphipods, small clams and hydroids, while polychaetes and barnacle nauplii may be found occasionally.
Lewis (1981) found some variability in stomach contents depending on the size of the P. polymerus. This was partly reflected by the change in method of feeding
with size (see p. 342). Three size ranges of barnacle were examined, 1–6, 7–14 and 15–28mm RC. The
Table 5 P. polymerus: food items identified in the midgut of various sizes of barnacle, n= number of animals examined; RC=rostrocarinal width. (After
Lewis 1981).
Percentage of barnacles in which food item was found Size range of barnacles, mm RC (n)
Food item 1–6 (9) 7–14 (17) 15–28 (17)
Organic particulates (total) 80 63 52
Inorganic particulates (total) 12 6 8
Large organic (total) 8 31 40
Organic particulates
Detritus 100 100 100
Diatoms 100 100 100
Other unicellular phytoplankton 0 >50 <50
Bluegreen algae 0 <50 <50
Inorganic particulates
Shell and sand >50 >50–100 >50–100
Sponge spicules few few few
Large organic
Crustaceans >50 100 100
Barnacle exuviae <50 100 100
Copepods 0 100 100
Polychaetes 0 >50 >50
Eggs 0 few <50
Hydroids 0 few <50
Molluscs 0 few few
Large algae 0 few few
Echinoids 0 0 few
Shrimps 0 0 few
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organic particulate food items (<10µm diameter) decreased from 80, to 63 to 52%, respectively, and the large organic material (10μm to 5000μm diameter) increased
from 8, to 31 to 40%, respectively. The inorganic particulate material (10µm diameter) remained virtually constant 12, 6 and 8%, respectively (Table 5). The
composition of the diet clearly indicates the change of feeding method between the three size ranges. The beating behaviour of the cirri was only seen in juvenile P.
polymerus and they mainly fed on small particulate material. Cirral beating may therefore be an adaptation for microphagous feeding. As has been shown, juveniles are,
however, capable of capturing larger food items. The beating behaviour of juveniles may be important in areas where there is no competition with the macrophagous
adults. The juveniles attached to peduncles of adults are below the mean cirral net level of the adults in a cluster.
The absence of sand in adult Pollicipes stomachs is probably because of the method of feeding in this species. As has been shown by Barnes & Reese (1959) the
capitorial action of the cirri was not readily elicited by inorganic material. The extra stimulus of a struggling prey enhanced the uptake of sand grains. There was little
evidence of any chemotactic sense involved as far as the cirri themselves were concerned. If sand grains are transferred to the mouthparts irritation of inorganic matter
may inhibit the cirral response. This species has a well defined rejection mechanism but this usually only removes finer material.
Allocation of energy
The effect of water temperature and food on the allocation of energy in P. polymerus has been examined experimentally by Page (1983). He used material collected
for the midintertidal level at Goleta Point, California. The animals used were 16.0 +1mm CH(±SD). He used Artemia salina adults, 4747+46calgDW−1 as food in his
feeding experiments. Two ration levels were used at 13.2±0.9°C and 19.8±0.5°C. Each day 2 (low ration) or 12 (high ration) Artemia per individual barnacle were
added to his experimental containers. The experiments ran for 40 days in May to July. His results are summarized in Table 6.
Table 6 P. polymerus: summary of number, dry weight, and calories ingested as Artemia and dry weight and calories egested as faeces in each
treatment. (After Page 1983).
Treatment
13.2 ± 0.9°C 19.8 ± 0.5°C
High ration Low ration High ration Low ration
Initial number of barnacles 50 50 50 50
Final number of barnacles 50 45 46 50
Ration (no. Artemia ind. −1 −1
day ) 12 2 12 2
Average no. Artemia ingested ind.−1 day−1 8.9 1.9 9.2 1.7
Total no. Artemia ingested ind.−1 treatment−1 356 76 368 68
Dry wt. Artemia ingested ind. −1

(mg) 126 30 129 23
Calories ingested ind. −1 598 142 611 109
Faeces, dry wt. ind.−1 (mg) 7.7 2.2 6.7 1.1
Faeces caloric content (cal g−1) 4518 4493 4934 4476
Calories egested ind. −1 34.8 9.9 33.1 5.1
Calories absorbed ind. −1

(ingestedegested) (A) 356 132 578 104
Absorption efficiency (%)
Dry weight 93.9 92.6 94.7 95.1
Calories 94.2 93.0 94.5 95.4
Ashratio 37.5 40.7 26.3 33.9
Ash absorption (%) 90.9 88.4 93.1 93.0
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Individuals on high ration ingested about five times as many calories as those on low ration. Absorption of ingested food (93–95%) measured by gravimetric
methods was not affected by temperature or ration. The ashratio method greatly underestimated the absorption efficiency. Weight of total body tissue only increased
in animals fed the high ration and at a higher temperature, but at high rations and low temperature the caloric content of both soma and stalk increased. There was a
decrease in the number of animals with egg lamellae, from 18 to 0%, and in the weight of ovary after 40 days in the warmwater cultures and on starvation. Food ration
did not have a significant effect on reproduction at either temperature. Welldeveloped ovaries have a caloric content of 5128 ± 15cal.gDW−1. P. polymerus embryos
take 25–30 days to develop (Hilgard 1960, Lewis 1975b) so the increase in number of animals with egg lamellae in the lower temperature cultures during the 40 days
suggests that these egg masses were produced sometime during the 40 days.
The energy values were obtained by multiplying the tissue dry weight by the corresponding caloric content (Table 7). These values (allowing for loss due to exuviae),
compared with the original sample, give the “actual” change in energy content of an individual during the 40 days. This is the production (P). The greater influence on
change in total energy was food rather than temperature. The increase in weight resulted in increased energy content at high ration and high temperature, whereas the
increase in energy in high ration and low temperature was the result of an increase in caloric content and production of ovarian tissue. The 20% decrease in weight of
starved animals accounted for their loss in energy content.
An estimate of the weightspecific respiratory rate at the end of 40 days is shown in Table 7. This rate increased in wellfed barnacles independently of an increase in
body weight. Two estimates of the absorbed energy used in respiration can be obtained from the general energy budget equation:
A=R+P+E,
where A=absorbed energy (calories),

R=energy used in respiration,
P=production measured as changes in energy content including moult and ovary production,
and E=excretion of soluble materials, not taken into account by Page (1983).
The first estimate (R=A−P) assumed a balanced energy budget. The second was the amount of oxygen consumed in each treatment multiplied by 4.8cal.mlO2−l.
This, divided by the absorbed calories gives the respiratory fraction (Table 7). The increase in respiratory rate of animals with the higher ration was probably related to
changes in their nutritional state. Temperature had little effect on the respiration of freshly collected P. polymerus (Q10=1.57).
The “estimated” energy available in each treatment for growth and reproduction is the socalled “scope for growth” (Table 7). High ration treatments at both
temperatures have a positive scope for growth, energy therefore being available for reproduction and growth. The negative scope for growth in the other treatments
indicated that, for maintenance, these animals must metabolize their body tissues. The estimated energy required for maintenance is 6.8cal. hid.−1 day−1. There was no
consistent temperature influence on the “actual” and “estimated” energy available for growth. There was good agreement between the two for animals fed the higher
ration and in starved animals. In animals fed the lower ration, the measured respiratory rate was higher than predicted from changes in energy content. P. polymerus is
intertidal and as such it is not surprising that food rather than environmental temperature influences production.
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Table 7 P. polymerus: average body weight (+SD), caloric content, reproductive condition, change in energy content (production), estimates of weight
specific respiratory rate, respiration fraction, and scope for growth of barnacles in each treatment. For values of A see Table 6. (After Page
1983).
Treatment
Initial 13.2+0.9°C 19.8±0.5°C Starved
High ration Low ration High ration Low ration
Body weight (mg dry wt)
Soma 56±14 56±11 51+13 53+12 45 ±16 48+8
Stalk 236 ± 57 232+63 214 ±52 267 ± 72 235 ±48 188 ± 40
Total 292 ±71 288 ± 84 265 ±73 320 ±60 280+62 236 ± 35
−1
Caloric content (calg )
Soma 4708 4819 4717 4684 4657 4694
Stalk 4633 4934 4881 4692 4655 4620
Exuviae
Average no. ind.−1 – 1.19 0.77 1.75 1.20 0.47
Reproductive condition
% brooding 18 25 17 0 0 0
Brood wt (mg dry wt) 20+6 19+5 14 ± 8 0 0 0
Ovary wt (mg dry wt) 4+7 10+5 6+4 1 ±2 1±1 1+2
Energy content (cal ind.−1)
Soma 264+67 270+53 241+59 248 ± 54 210+73 225 ± 37
Stalk 1093+359 1145+359 1044+386 1253 ± 360 1094 ±430 870+198
Ovaries and brood 21 51 31 5 5 5
Total 1378 1466 1316 1506 1309 1100
Exuviae – 21 13 28 21 7
Production (P) (Δcalind. ) −1 – +102 −48 +156 −48 −272
Oxygen consumption rate (ml O2gDW−1 h−1)
Day 0 – 0.24 0.23 0.34 0.32 0.24

Day 40 – 0.44 0.24 0.42 0.32 0.25
Respiration fraction
Estimate from production (1P/A) – 0.81 1.36 0.73 1.46 –
Estimate from O2 consumption (R/A) – 0.73 2.42 0.88 3.96 –
Estimated calories expended respiration from O2 consumption measurements (R) – 411 319 509 412 314
Scope for growth (AR) – +152 −187 +69 −308 −314
Effect of temperature change and desiccation

As with other intertidal cirripedes, Pollicipes species have to tolerate stresses due to temperature change and desiccation when exposed to air during intertidal periods.
Pollicipes has a wide range of distribution and must, therefore, have a high tolerance to adverse conditions. It has been found that P. polymerus can withstand 5–10h
exposure to air at 36°C (Fyhn et al.
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1972) as well as at 2°C (Petersen et al. 1974). The wide distribution of P. polymerus suggests that it is a eurythermal species. When submerged the animal assumes
the seawater temperature but when in air, there are appreciable differences between body temperature and the ambient. The effect of sun is to raise the body
temperature well above air temperature and it stabilizes at 2–5°C below the temperature of the substratum to which the animal is attached.
When insolation was prevented in experimental conditions the body temperature of P. polymerus was found to be lower than air temperature and the lowering of
temperature increased with increasing air temperature. At 36°C and 55% relative humidity capitulum and peduncle temperatures were 4 and 8°C, respectively, below
air temperature. This cooling by evaporation may have survival value by increasing the heat load an animal can tolerate. Heat loss in Pollicipes seems to be mainly from
the peduncle due to the water evaporating there. Water loss from the capitulum was almost negligible and temperature depression in the capitulum may be an effect of
cooled haemolymph circulating from the peduncle and not due to evaporation (Fyhn et al. 1973).
The rate of loss of water by evaporation from the cuticle of the peduncle of P. polymerus was about 1.5μlcm−2h−1mmHg at 21°C and 27°C showing that this
barnacle is more permeable to water than many balanids. Experimental animals were shown by Fyhn et al. (1973) to recover after 40–50% water loss. These authors
calculated that at the above rate of water loss a Pollicipes of 25 g weight and an area of peduncle surface of 35cm2 would lose 0.75 ml water h−1 at 27°C and 55%
relative humidity. With an animal water content of about 77% this amounts to a loss of about 4% body water per hour. The time Pollicipes is exposed between tides is
less than 9h and so animals will not lose more than 35–40% of their body weight during normal periods of exposure to air. This is within the tolerance limits. A water
loss of 0.75mlh−1 is equivalent to heat loss of about 450 cal.h−1 (Fyhn et al. 1972). Water lost from P. polymerus during an intertidal period has to be replaced during
the next submersion. In experiments it was found that after exposure to air at 27°C and 55% relative humidity for 8h, 80% of the weight lost was recovered within 4h
when resubmerged. The initial rate of rehydration was rapid but after about 4h it slowed down and after 10–14h the initial weight was reached. Rehydration at high
tide levels has to be rapid as time of resubmersion is less than lower on the shore.
Respiration, gaseous exchange and oxygen uptake

There is no specialized organ for respiration in pedunculates; they rely on gaseous diffusion across parts of their anatomy such as cirri, body surface, and mantle cavity
lining. There may also be some exchange across the peduncular wall especially in species with a relatively thin cuticle. In the three species of Pollicipes there are
filamentary appendages at the base of the first cirri. These and the prosomal filamentary appendages present in these species may act as branchiae (Walker 1992).
There is, therefore, a considerable surface area exposed to the water current ventilating the mantle cavity. Ventilation is dependent on cirral activity and it is sufficient for
normal oxygen requirements. According to Anderson & Southward (1987) and Anderson (1994) the amount of ventilation may be regulated by a receptor that is
sensitive to carbon dioxide concentration.
Barnes & Barnes (1959a) used excised bodies to examine the oxygen uptake of a variety of cirripedes in an attempt to relate it to species habitat. It was found that
generally the intrinsic metabolism decreased from sublittoral to high intertidal species. It is curious that P. polymerus
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was nearer to the balanids than to the Chthamalus species tested. Even though excised bodies survived for many hours Petersen et al. (1974) thought using them might
give misleading results. Despite this, when Page (1983) converted the results of Barnes & Barnes (1959a) to the same units as his so that comparisons could be made,
he found a good agreement.
In an intertidal cirripede gaseous exchange has to take place in water and while exposed to air. In Pollicipes polymerus there is actually an increase of aerobic
metabolism during emersion (Petersen et al. 1974) in contrast to balanomorph barnacles which have a reduced rate of oxygen uptake during aerial exposure. This could
be associated with the rather sluggish cirral activity in Pollicipes so that cessation on exposure to air does not make much difference to the respiratory needs. Cirral
activity in balanomorphs is far more energetic. An adequate surface for gaseous exchange must be available if the oxygen uptake is to remain unchanged on emersion.
As the opercular valves close when the cirri are withdrawn on exposure to air the surfaces within the mantle cavity have to be amended. There are haemolymph sinuses
below the thin cuticle of the mantle cavity and Burnett (1972) suggested that these might be involved in gas exchange. Petersen et al. (1974) suggests that there may be
additional gas exchange across the wall of the peduncle. For this to be effective an efficient haemolymph circulation in the peduncle is important. Such a high pressure
circulation has been found in P. polymerus by Fyhn et al. (1973).
Petersen et al. (1974) found that oxygen uptake in P. polymerus increased with temperature both for animals exposed to air and when submerged. In the former
conditions the animals had a respiratory rate five times higher at 10°C and three times higher at 20°C than when submerged. Submerged animals had a Q10 of 2.5
between 10°C and 20°C, while in air it was 1.6 between 10°C and 20°C. In air there is a decreased dependency on increasing temperature: Q10 of 2.2 between 2°C
and 10°C reducing to 1.6 between 10°C and 20°C, and 1.3 between 20°C and 27°C. P. polymerus was found to show oxygen conformity under varying ambient
oxygen tensions; no oxygen debt was found after exposure to hypoxic water.
Innes (1985) examined the oxygen uptake and haemolymph oxygen tensions in Calantica spinosa (=Pollicipes spinosus) and found that the aerobic respiratory
system functioned equally well in water and when exposed to air. Petersen et al. (1974) found that in P. polymerus the aerobic metabolism was also positively
enhanced in air. It has been suggested that this may be a general feature of intertidal pedunculate barnacles and probably contributes to their success in the intertidal
community.
The effect of temperature on the oxygen uptake and rate of development of eggs of P. polymerus was examined by Barnes & Barnes (1959b). Samples were taken
from Friday Harbor, Washington State, and Santa Monica, California—some 1000 miles (latitudinal) apart. Irrespective of the stage of development of the embryo or
the temperature at which the estimations were made, there was no indication of any regional difference in oxygen uptake (Table 8, values from 15°C to 25°C). In P.
polymerus there is a marked change at 10°C in all except the very young stages of development. The oxygen uptake of the younger stages is
Table 8 P. polymerus: oxygen uptake (μl g−1 h−1) of egg lamellae at different stages of development and different temperatures. SM=Santa Monica,
California; FH= Friday Harbor, San Juan Island.(After Barnes & Barnes 19595).
15°C 20°C 25 °C
Stage SM FH SM FH SM FH
Early Stages – – – – 237 245
Middle Stages – – 342 323 375 410
Ready to hatch 190 188 282 305 – –
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relatively independent of temperature, Q10 4.2. Beyond this stage and above 10°C the Q10 approximates to 2.5. When corrected for the difference in nitrogen content
it is of interest to note that the rate of oxygen uptake, on the basis of a single egg, in P. polymerus approximates to onesixth to onethird that of a Semibalanus
balanoides egg. As far as can be seen from the results there is no acclimation as regards oxygen uptake in the embryos of Pollicipes polymerus.
Chemical composition
Bodies and egg lamellae

The distribution and fatty acid composition of the main lipid class in ovary, egg lamellae, and bodies (prosoma and thorax) of several cirripede species including P.
pollicipes have been determined by Morris & Barnes (1975). P. pollicipes was collected by divers from an exposed situation typical for this species at Biarritz,
France. The egg lamellae and body were removed from the mantle cavity and the ovary was dissected from the peduncle. Details of the methods of analysis are given
by Morris & Barnes (1975) and not repeated here. The relative amounts of the lipid classes, the fatty acid composition of the triglyceride (TG) and phospholipid (PL)
and the relative distribution of mono and polyunsaturated acids are given in Tables 9, 10 and 11.
The egg lamella tested were pooled samples and contained embryos at all stages of development. A newlyfertilized egg will be little different in composition from the
ovary so the greater PL must be a consequence of the development of the embryos, such as the development of membranes. Both TG and PL fractions in the egg
lamellae contain a higher proportion of polyunsaturated fatty acids than the ovary. Because the lamellae are metabolically “isolated” such changes must represent
synthetic processes in the developing embryos. This increase in polyunsaturated fatty acids cannot be an adaptation to low temperature because breeding and embryo
development in a warm water species such as P. pollicipes is in the warmest months. This may not apply in P. polymerus because of its wider distribution on the
Pacific coast of North America. There are changes in phosphorus metabolism of embryos during their development. The increase in polyunsaturated fatty acids is due
to increased formation of membranes resulting in additional structural PL.
The difference between the fatty acid composition of body and ovary does not support the suggestion that ovary TG is derived directly by transfer from the body—
unless further changes take place after transference. The ratios of 20:5/22:6 fatty acids indicate that these two acids are nearly equal or at least more nearly so in all
components than the results found in some balanid species given by Morris & Barnes (1975).
Table 9 P. pollicipes: lipid, triglyceride (TG), and phospholipid (PL) composition. (From Morris & Barnes 1975).
Sample Lipid, est. % wet wt. TG, % total lipid PL, % total lipid
Ovary 2.93 60.9 21.7
Egg lamellae 1.98 64.2 30.0
Body 1.02 43.0 40.0
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Table 10 P. pollicipes: fatty acid composition of lipids (% total fatty acid). T=trace <0.5% (limit of detection 0.1–0.2%); * may include other isomers.
PL=phospholipid; TG=triglyceride. (From Morris & Barnes 1975).
Doublebond Ovary Bodies Egg lamellae

Fatty acid position PL TG PL TG PL TG
12:0 T 1.1 – T T T
iso 14:0 – – – – – –
14:0 1.6 7.0 2.1 6.7 1.3 3.3
14:1 5* T 0.5 T T T T
ante 15:0 – – – – – –
15:0 T 0.8 T 0.5 T 0.5
iso 16:0 T T T T T –
16:0 17.9 21.5 24.1 29.2 19.3 20.7
16:1 7* 2.9 7.0 2.5 9.7 2.6 6.1
16:2 9, 12 – T – – – –
17:0 0.9 0.9 1.2 0.6 1.0 T
17:1 ? T T 0.5 0.8 T T
iso 18:0 T T T T T T
18:0 11.4 5.5 13.0 5.2 11.1 6.2
18:1 9* 18.2 18.6 13.4 21.4 15.3 20.8
18:2 9, 12 3.4 4.8 2.8 3.4 3.2 5.0
18:3 6, 9, 12 T T – T T T
18:3 9, 12, 15 0.5 0.8 T 0.8 0.5 1.0
18:4 6, 9, 12, 15 0.6 0.6 5.8 0.6 0.6 0.6
19:0 T T – T T –
iso 20:0 – – – – – –
20:1 11* 3.3 3.3 3.0 2.6 3.4 3.2
20:2 11, 14 T – T 0.8 T –
20:4 5, 8, 11, 14 2.8 0.6 1.3 0.7 1.3 0.8
20:5 5, 8, 11, 14, 17 19.2 10.8 13.9 9.0 20.2 13.3
22:1 13* T 0.5 0.6 0.6 T 0.5
22:3 7, 10, 13 – – – – – –
22:3 7 – T – T – T
22:4 10, 13, 16, 19 – T – – – T
22:5 7, 10, 13, 16, 19 1.2 T 1.4 T T T
22:6 4, 7, 10, 13, 16, 19 14.1 15.5 12.8 7.2 18.5 17.3
Ratio 20:5/22:6 1.4 0.7 1.1 1.2 1.1 0.8
Changes in the general biochemical composition of the developing eggs of P. pollicipes have been determined by Achituv & Barnes (1978). The egg lamellae were
separated into three groups; the eggs in a single lamella are all at approximately the same stage of development. Pooled samples were used. The groups were: Group 1,
early from first division to few cells; Group 2, multicellular; and Group 3, nauplius eye visible. The results are given in Tables 12 and 13.
The energy content per unit volume of egg material, that might be called the related volumespecific caloric content, is obtained by dividing the energy content per
106 eggs by the egg volume. In this way the result obtained for P. pollicipes (Table 13) can be compared
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Table 11 P. pollicipes: relative distribution of saturated mono and polyunsaturated fatty acids in lipids (% wt of total fatty acid). (From Morris &
Barnes 1975).
Ovary Egg lamellae Body

Triglyceride
Saturated 37.0 31.4 42.2
Monounsaturated 30.2 30.6 35.1
Polyunsaturated 33.1 38.0 22.5
Phospholipid
Saturated 32.2 33.4 41.1
Monounsaturated 25.1 22.0 20.0
Polyunsaturated 42.2 44.7 38.6
Table 12 P. pollicipes: biochemical composition of embryos (mg 10−6 eggs) and cumulative percentage change during development. (After Achituv &
Barnes 1978).
Group 1 Group 2 Group 3

Dry weight 657 547 530
Cum. loss – 16.7 19.3
Carbohydrate 31 26 14
Cum. loss – 16.1 52.9
Protein 321 292 269
Cum. loss – 9.0 16.2
Lipid 178 108 93
Cum. loss – 39.3 47.8
RNA 7.6 10.6 13.9
Cum. gain – 39.5 82.9
DNA 2.0 – 3.6
Cum. gain – – 44.4
Ratio protein/lipid (P:L), wt/wt 1.80 – 2.88
Table 13 P. pollicipes: energy values during development. Conversion factors used: carbohydrate, 4.1 kcal g−1; protein, 5.65 kcal g−1; lipids, 9.45 kcal
g−1. (After Achituv & Barnes 1978).
Energy content,
initial, kcal 10−6 eggs 3.62
final, kcal 10−6 eggs 2.46
Egg volume (V)×10 −6

ml 1.41
Caloric value, % lost 32.00
Ratio initial caloric values, kcal/V 2.57
Ratio final caloric value, kcal/V 1.74
with other results published by Achituv & Barnes (1978), Pollicipes has a value (2.57) similar to that of Tetraclita squamosa rufotincta (2.68) and Chthamalus
stellatus (probably C. montagui) (2.65). Values for some balanid species were considerably lower.
Two of the substrates used during the development of eggs are protein (p) and lipid (L). The ratios of these (P:L) at the beginning and end of development are given
in Table 12. The utilization of lipid in preference to protein was evident in Pollicipes and the final ratio (2.88)
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was the highest of all the other cirripedes (1.83–2.38) quoted by Achituv & Barnes (1978). Of all the cirripedes examined utilization of lipid in Chthamalus was
nearest to that used in Pollicipes, but use of protein was least in Pollicipes. Increase in DNA reflects tissue development and organization; RNA is associated with
protein metabolism and continues to increase during development. Holland (1987) has reviewed the lipid biochemistry of barnacles and mentions the relationship
between carbohydrate, protein, and lipid in the development of egg lamellae.
Calcified structures
Only in relatively recent years have studies been carried out on the internal structure and composition of the barnacle shell. Bourget (1987) has reviewed the present
position and gives all the relevant literature. Even now there is very little information on pedunculate cirripedes.
The two major components of a barnacle shell are calcium carbonate (in the majority of cases this is calcite) and organic matter. The overall organic content and the
ratio of calcium to magnesium have been determined for several species including Lepas fascicularis and Pollicipes polymerus by Barnes et al. (1976). From Table
14 it can be seen that P. polymerus has lower values than Lepas for total organic matter, protein, and chitin; in fact the values were lower than those found in any other
species tested. 35SO4 incorporation into Pollicipes polymerus also indicated a low organic component in the calcite (Chaffee & Lewis 1988). A marked correlation
was found between the calcium:magnesium ratio and the organic matter in all the species examined including the Lepas species. As changes in organic matter reflect
changes in protein content this supports the suggestion that magnesium may be associated with organic matter rather than the calcite. Values for Pollicipes polymerus
were anomalous, having a much lower organic content than would have been expected from their calcium : magnesium ratio; they were outside the correlation. Koehler
(1889a) found that the structure of the calcified tissue in Pollicipes was more complicated than in other pedunculates.
Table 14 Composition of shell plates (from Barnes et al. 1976)
Pollicipes polymerus Lepas fascicularis

Total organic matter, % 0.39 0.83
Protein (p), % 0.31 0.72
Chitin (c), % 0.08 0.11
Ratio, P/C 3.88 6.55
Calcium: magnesium ratio 47.8 74.0
Moulting and growth

The connection between moulting and growth in cirripedes has been well summarized by Anderson (1994). He clearly states the difference between the processes in
pedunculates and balanids and quotes the appropriate literature. Stalked barnacles such as Pollicipes species grow in height by increasing the length of the peduncle
and in width of capitulum by addition to the calcified plates by accretion.
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The narrow zone between capitulum and peduncle in stalked barnacles has been called the “summit of peduncle” (Darwin 1851), “transition zone” (Broch 1921,
1922), and “zone génératrice” (Prenant 1924); Mahmoud (1959b) decided to call it “growth zone” and this term has persisted. It has long been recognized that this is
the area where new scales are developed as reported by Foster (1978). He found that plate multiplication occurs in P. polymerus at the base of the capitulum. In this
way those already on the capitulum are elevated towards its apex. The number of whorls of these plates depend on the age of the animal. Darwin (1851) found two or
more in all three Pollicipes species.
On the capitulum each primary plate (scuta, terga and carina) originates as a primordium (Broch 1922) that then becomes a point of calcareous accretion (see
Anderson 1994; see Fig. 4, p. 320). Areal growth is by marginal accretion. Each plate has an underlying epidermis and grows in thickness by laminar accretion (see
also Koehler 1889a) over the whole inner surface. Secondary capitular plates that develop below the primary plates are added to in whorls from the growth zone,
those closest to the zone being the youngest. In Pollicipes there may be several additional whorls of smaller and smaller sublateral plates at the base of the capitulum.
Secondary plates begin directly as calcareous structures as do the peduncular scales. Secondary capitular plates retain only basal contact with the underlying epidermis.
Accretion is mainly basal, extending on to the basal margins of the inner and outer surfaces so that additions are concentric. The rostrum, rostrolaterals, carinolaterals,
and subcarina are all more closely adpressed to the basal junction of scutum and tergum on each side and therefore there may be more accretion on the inner side
allowing some laminar growth. As they grow the sublateral plates project more than the lateral ones.
Some small calcareous scales produced in the growth zone are displaced on to the lengthening peduncle (Fig. 14). Darwin (1851) counted 10 whorls of developing
scales in the growth zone of P. pollicipes, but the numbers may vary from 6 to 8 in young animals and 8 to 12 on older ones (Mahmoud 1959b). Broch (1922) found
that the appearance of new peduncular scales in the growth zone of P. polymerus continued throughout the animal’s life.
It is usually accepted that the outer integument of the capitulum and peduncle is not shed at a moult as is the inner integument. Even so Mahmoud (1960), when
studying P. pollicipes, described what he calls normal and regular moulting of the cuticular layer of both the capitulum and the growth zone, but there was no moulting
of the peduncle proper. The moult, therefore, is occurring in regions where the cuticular formations (plates and young scales) are
Figure 14 Pollicipes pollicipes: diagramatic representation showing continuous development of new scales in growth zone (A to E)
and the increase in length of the peduncle after moulting of the growth zone (B and D). Note the progress of scales (1–8)
down the peduncle. C=capitulum; GZ=growth zone; m=moult; P=peduncle. (After Mahmoud 1960).
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attached to the ectoderm. Moulting of the capitulum, however, was only found in young animals but continued throughout life in the growth zone. This allows the
appearance and development of new scales.
A splitting of the old cuticular layer, between the different plates and developing scales, from the underlying ectoderm and, at the same time, the formation of a new
cuticular layer covered externally by epicuticle initiates the beginning of moulting on either the capitulum or in the growth zone. Simultaneously an exuvial epicuticular
lamina appears separating the lower limit of the growth zone from the peduncle proper. If only the growth zone is moulting a similar lamina separates the upper limit of
the growth zone from the capitulum. The thickness of the new cuticle increases from its inner surface and is pushed out over the plates and developing scales until at the
end of the moulting processes the upper young scales are covered with a solid cuticular layer. At the same time, the old cuticle gradually breaks down.
This “moulting” described by Mahmoud (1960) is not true moulting as occurs in the lining of the mantle cavity and the soft parts (thorax, cirri and prosoma). It is
probably better described as a roughening of the cuticle that becomes scaley and pieces of which break away from the surface.
When first formed, a peduncular scale is in the uppermost whorl in the growth zone and when fully formed it is in the uppermost whorl of the peduncular scales
proper. Successive stages of development are found in between these two whorls. The apparent “migration” of scales is caused by an extension in length of the
peduncle from its uppermost part (see Fig. 14).
Radioisotopes have been used in the most recent study of the growth of the exoskeleton and penduncular scales of P. polymerus (Chaffee & Lewis 1988). 3H
thymidine was used as a marker of mitotic activity and was found to concentrate in the growth zone and at the base of the peduncle. New exoskeleton for the peduncle
is formed in the growth zone and is continuous with the earlier exoskeleton (Darwin 1851, Mahmoud 1959a, 1960). The latter author found that the new exoskeleton
could not be distinguished from the older one. When grown in low light conditions and fed differently coloured food the coloured pigment was found in cuticular bands
in the growth zone. Such bands are difficult to see in animals grown in normal light conditions because of the animal’s own pigmentation that darkens to purplishblack
(Barnes & Reese 1960). The feeding experiments done by Chaffee & Lewis (1988) showed that peduncular growth was intermittent and the number of new bands
was approximately equal to the number of body exuvia recovered from the culture dishes. It cannot be said, however, that band formation and body moulting always
occur simultaneously.
Chaffee & Lewis (1988) found that the band width of exoskeleton ranged from 1–3 mm in 9 days, but could be wider in animals from different environments. Young
animals grown in strong currents with plenty of food could produce bands of 10mm width and grew 100mm in peduncle length in 2–3 months (see also Hoffman 1988).
Growth zones in P. pollicipes have been confirmed by Williams (1991).
Different rates of calcification in peduncular scales compared with capitular plates were found by Chaffee & Lewis (1988). Mahmoud (1959b) regarded calcification
as a continuous process throughout the life of P. pollicipes. Chaffee & Lewis (1988) agreed but believed rates of calcification varied according to environmental
conditions. They considered it possible that calcification and stalk growth could decrease or even stop under stress conditions such as lack of food or desiccation due
to high temperatures.
The mitoticallyactive site found by Chaffee & Lewis (1988) at the base of the peduncle explains observations made by several other workers. This growth was
particularly noticeable in young animals but was also found in adults within clusters as a bulbous area at the
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Figure 15 Pollicipes polymerus: juveniles settled on an adult peduncle. Note absence of Pollicipes on the capitulum of the adult on
which there are several adult Chthamalus fissus.
base of the peduncle. Cyprids and young spat are almost always found on peduncles of older Pollicipes as this provides an ideal substratum for settlement (Fig. 15). A
clump of older barnacles provides cyprids with vacant sites and instant protection from physical elements among the numerous peduncular scales. The need for light and
chemical recognition attracts the cyprids to the growth zone. Hoffman (1981, 1984, 1989) found that in time young P. polymerus were able to detach themselves from
the adult peduncle, presumably regarding it as only a secondary and nonpreferential site. By twisting their peduncles round adjacent peduncles the young animals were
held in place while they extended their peduncles downwards eventually reattaching on the primary substratum. T. Cruz (pers. comm.) found the same phenomenon in
P. pollicipes and this has been confirmed by Kugele & Yule (1993). These two found evidence of tracks of animals that increased in width with increasing distance
from the beginning of the track thus indicating the direction of movement. Interaction between muscular activity and growth may cause the mobility. Haemolymph
pressure could distort the end of the peduncle in the direction of travel. These extensions often appear bright red because of the thinness of the cuticle and the
haemoglobin contained in them (Hoffman 1984). Reduced growth of the trailing region and muscular activity needed to shear the old adhesive could maintain direction.
The downward direction of juveniles on the adult barnacle could be the result of a positive reaction to gravity and a negative one to light. When adults
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are turned through 180° the juveniles changed direction towards the capitulum, but still downwards. The movement of two juveniles of rostrocarinal length 6.9mm and
8.3mm, respectively, in a seawater flow of about 8cms−1 was approximately 0.06–0.08mmday−1, measured over 55 days. Kugele & Yule (1993) maintained that the
movement of young adults resulted in a more even distribution of similarsized young on adult peduncles than that found be Hoffman (1984) for P. polymerus.
Movement of juveniles from the upper part of adult peduncles, eventually reaching the primary substratum, is advantageous in preventing overcrowding and
smothering of adults in the upper layers of the clusters, while still maintaining the cluster formation so characteristic of the species. This movement of juveniles involves a
reattachment to the primary substratum when it is reached; for this a secondary secretion of cement must be produced (Hoffman 1984). Meese (1993) made several
attempts to transplant Pollicipes for experimental purposes but was not successful, nor was he successful in getting any settlement on plates. Williams (1991) found,
however, that unattached P. pollicipes could produce secondary cement and she found two animals in experimental conditions that had become attached to each
other.
The settlement period begins when an animal first lands on a substratum, that is when a cyprid has cemented itself to the surface. Recruitment is a measure of
recentlysettled juveniles that have survived for a period of time after settlement. There have been very few studies that compare settlement and recruitment in
cirripedes; the cyprids settle in cryptic habitats such as on other organisms or in crevices. During the first 2–4 wk after settling they are subject to great mortality.
The cyprids of Pollicipes settle on peduncles of conspecifics and so settlement, recruitment, and juvenile growth rates can be determined with relative ease. Hoffman
(1989) investigated settlement in a variety of habitats on the coast of California near La Jolla. Settlement of P. polymerus cyprids occurred all the year, the peak being
in early spring when he found between 150 and 300 spat ( ≤1mm RC) per adult peduncle. He only found spat on the primary substratum on the under surface of a
boulder that was presumably continually washed by waves. Peaks of survival of juveniles were during summer, which related to the highest spat peak in spring.
Recruitment occurred throughout the year. Juveniles between 1 and 9mm RC remained on the adult peduncles but they produced extensions at the base of their
peduncles and, after movement downwards, cement from the swollen ends eventually enabled them to reattach themselves to the primary substratum (see above).
Some of these formed new clusters and growth in them was rapid. When continually submerged, juveniles could reach 9mm RC in about 1 month. On vertical surfaces
it could be 7 months before juveniles became established. Hoffman (1989) found survival from spat to juveniles was highest on solitary individuals and lowest when
mixed with mussels. The low midintertidal level seemed to favour the best recruitment.
Size distributions of juvenile P. polymerus on adult peduncles have been made by Hoffman (1984) on material from the coast of Washington State and from
Monterey Bay, California. He found that the young animals were not randomly distributed. There was a gradient from the smallest and most abundant near the growth
zone to the largest and fewest located near the base of the peduncle. It was extremely rare to find a cyprid or young spat on the capitulum of an adult although
Chthamalus species or Balanus glandula may be found there (see Fig. 15, p. 356). Broch (1928) found one cyprid crawling on the capitulum of Pollicipes
pollicipes. When the adults were constantly immersed in running sea water cyprids have been found to settle on the capitular plates and valves and to grow into
juveniles (Hoffman 1988). This is an indication that in natural intertidal conditions the adult peduncle may provide better protection from desiccation than the capitulum.
The smallest juveniles of
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P. polymerus found among the peduncular scales were 0.5mm in total contracted length.
In experiments done in a seawater intake flume 5m from the seaward end Pollicipes larvae settled in great numbers (Hoffman 1988). Cyprids were also found to
settle on roughened areas of terracotta tiles but not on mussel shells nor on balanids. Within two weeks of settlement on the tiles the characteristic aggregates of spat
were evident due to the presence of conspecifics. Lewis (1975b) did not observe settlement of Pollicipes cyprids in beakers of aerated sea water; there was no
settlement on etched glass slides, nor on plates coated with extracts of adult barnacles, nor on natural rocks from the field although Lewis was aware that a combination
of chemical and tactile responses was necessary for the specific settling behaviour of P. polymerus. In the flume, where constantly immersed and at a temperature of
20°C, Hoffman (1988) found juveniles had a growth rate of 5.49mm RC in 47 days with a range up to 11 mm. Such a rapid growth rate for some other pedunculates
was also found on continually submerged buoys by MacIntyre (1966).
Bernard (1988), in connection with the development of potential fisheries for P. polymerus, studied the recolonization of denuded areas of the coast of Vancouver
Island, British Columbia. He reported settlement of cyprids as occurring “indiscriminatly on solid substrates [sic] from the upper intertidal to 3 [m?] below the lowest
tidal level”. Such settlement in the upper intertidal is at variance with that found by Hoffman (1989) but it may be caused by an initial settlement of Balanus glandula on
the denuded areas examined by Bernard (1988). He does say that the settlement of Pollicipes polymerus depends on an initial settlement of the balanid as this
provides some early protection for the spat. It may also be that these “denuded” areas had previously supported P. polymerus populations; there may have been some
residual conspecific attractant on the surface. Bernard (1958) found settlement from March with peaks in June and July. Early growth was rapid and comparable with
other results (see Table 15).
The effect of settlement density on the spatial arrangement of intertidal barnacles has been examined by Satchell & Farrell (1993). The degree of aggregation in P.
polymerus increased significantly as density of settlement increased. This is what would have been expected from earlier work (Lewis 1975b, Hoffman 1984, 1988,
1989). The disadvantages of an aggregated intraspecific life style are probably outweighed by the reduction of desiccation of newlysettled cyprids, the influence on
water flow to enhance feeding possibilities, provision of additional support to the developing peduncles of young animals as they relocate after settlement on the primary
substratum, exclusion of interspecific competitors (Hui & Moyse 1987), and protection from predators such as birds (Meese 1993).
Differences in population structure and growth rates were found in P. polymerus growing on a rocky headland, Goleta Point, Santa Barbara, California and on an
18yearold oil platform 3km offshore (Page 1986). There were no differences in the size structure of the populations but the structure was skewed at Goleta Point
with no animals >26mm CH and peaked at 6mm CH. The offshore population had peaks at 10 and 26–30mm CH and a maximum barnacle size of 38mm CH. A
single sample from another oil platform had peaks at 10 and 30mm CH and a maximum size of 34mm CH. The growth rate of individuals of 10mm CH was
0.114mmday−1 at the oil platform and 0.033mmday−1 at Goleta Point. The expected capitulum heights after 1 and 5 yr are given in Table 15.
It was found that only 8 % of the population at Goleta Point was >2 yr old whereas on the oil platform it was 30%. The decrease in number of individuals with
increase in size indicated that mortality was size and/or age dependent. The unstable shale substratum which tended to break away caused mortality at Goleta Point.
Older animals here tended to grow in bivalve holes or crevices and were protected from erosion. Younger animals settled on the
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Table 15 Growth rates of P. pollicipes and P.polymerus. RC is width across capitulum from rostrum to carina; CH is vertical height from base of
sublateral plates to tip of tergum; 1 RC approximately 0.84 CH.
Location Notes Growth rate Reference

P. pollicipes
SW coast, Portugal Ambient intertidal conditions Juvenile, 0.17–0.66 mm RC month−1 Adults, 3mm RC year−1 Cruz 1993
P. polymerus
Amphitrite Point, Vancouver Ambient intertidal conditions on cleared Juveniles, 4–5 mm RC in 5 months, reached 15 mm in first year Bernard 1988
Island, BC plots
Adults, possibly 26 mm RC by end of 4 years
Pacific coast of USA Ambient intertidal conditions Adults >17 mm, 0.5–0.7 mm RC in 16 months. >30 mm RC may be 20 years Barnes &
old, but no way of determining age. Reese 1960
Pacific coast of USA Adults slow growth after >10 mm—maturing in 5 years. Newman &
Abbott 1980
San Juan Island, Washington Ambient temperature 6.2–13.6°C Juveniles, 0.2–1.0 mm RC month−1 reached 15 mm in first year. Adults of Lewis & Chia
State depending on season, intertidal natural 1981
conditions RC=13mm, 1–2mm RC yr−1. Adults of RC approx. 30 mm, growth very slow
San Juan Island, Washington Ambient temperature 6.2–13.6°C Juveniles, 0.7–1.2 mm RC month−1 reached 18 mm in first year. Adults, 4.2 Paine 1974
State depending on season, intertidal natural
conditions mm RC yr−1 reached 35 mm in 6 yr
Goleta Point, Santa Barbara, Ambient intertidal conditions on unstable Juveniles, 7–9 mm CH (approx. 8.3–10.1 mm RC) in 10 months at 10 mm CH Page 1986
California substratum rate 0.033 mm d−1 15 mm CH (approx. 18 mm RC) in first year. Adults 20 mm
CH (approx. 23 mm RC) after 5 yr
Oil plaform, off Santa Intertidal conditions but 3 km offshore on Juveniles at 10 mm CH, rate 0.114 mm d−1, approx. 25 mm CH (approx. 30 Page 1986
Barbara stable substratum mm RC) in first year Adults approx. 34 mm CH (approx. 40 mm RC) after 5 yr
Scripps Insitution of Intake seawater system, 20°C, total Juveniles, 5.49 mm RC in 47 days including a max. of 11 mm 47 days Hoffman 1988
Oceanography, La Jolla, submersion in good water flow 5 m from
California seaward end
Coast near Scripps Under surface of rock continually in Juveniles, 9 mm RC in one month Hoffman 1989
contact with sea water
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new areas exposed when the shale broke away, thus increasing the proportion of younger animals. The metal supports of the old platform provide a stable substratum
for the animals.
On an experimental plot where the larger barnacles (>20mm CH) were removed Page (1986) found that after 98 days the barnacles were significantly larger than
their initial size and the size of the smaller ones in a control plot. The increase in size of 4mm CH (from 10 to 14 CH) was about half that predicted from the growth
curve (10mm CH to increase 4mm in 45 days). The stable substratum of the oil platform allows large clusters of barnacles to become established and intraspecific
competition probably restricts growth of recent recruits and most may not attain 10mm CH. Once the larger animals are removed the smaller ones have the ability to
grow. On the experimental plot this growth may still be inhibited by the larger animals remaining at the margins of the plots owing to their interference with the water
flow and availability of food for the smaller animals on the plot. Hence the slower growth rate than predicted for these smaller animals.
Factors affecting growth of P. polymerus have been mentioned several times and in order to grow the barnacle must moult. There is not much information on the
effect of food and water temperature on moulting in P. polymerus. Page (1983) examined moulting frequency of the species from Goleta Point. Ambient water
temperature at the time of collecting the experimental animals was 15.5°C. The capitulum height of the animals used was 16 ± 1mm (standard deviation). The
experimental chambers were whirlpools that provided the correct water movement for feeding. Four conditions with 50 animals in each were investigated in ambient
light conditions: low temperature, 13.2 ± 0.9°C, and high temperature, 19.8 ± 0.5°C each with high and low food ration. A control group of 15 animals was kept at
13°C and starved. The moulting frequency in each treatment was calculated as the average number of moults produced per day for each consecutive 10day period.
Water temperature and food had no effect on moulting during the first and second 10day periods. There was a significant interaction with temperature and food
between 21 and 30 days, whereas between 31 and 40 days food alone was responsible for influencing moulting. The high moulting rate at high temperature and low
food ration between 21 and 30 days followed by a drop between 31 and 40 days compared with a continued increase at high temperature and high food ration
suggests that high temperature may accelerate moulting and deplete food reserves after 30 days. The greater influence of food rather than temperature is probably to be
expected in this intertidal pedunculate. The availability of food when totally submerged has been seen to be an advantage for growth in P. polymerus (Hoffman 1988).
Preliminary results of factors influencing growth of P. pollicipes on the southwestern coast of Portugal have been given by Cruz (1993). Three sites representing
different variables were examined. Site 1 was from the midpart of a vertical distribution of the animal, +1.5m to 3m above MLWS (mean low water spring level). Site
2 was in the lower intertidal, MLWS to+1.5m. Site 3 was a shaded region in the middle of the vertical distribution. Several measurements were made including the
rostrocarinal width, capitulum height and peduncular length. Her results suggest that light could be an external factor affecting longitudinal growth of the whole animal.
The capitulum height and peduncle length were greater at Sites 2 and 3 than at Site 1. Body weight was greater at Site 2 than at Sites 1 or 3, probably reflecting the
greater immersion time and hence more available food. Rostrocarinal widths were the same at all three sites. Animals from different sites with the same rostrocarinal
width had the same peduncle weight. P. pollicipes was found to grow faster in spring and summer, almost certainly owing to the more abundant food and higher
temperature in these seasons. Growth rates of juveniles ranged from 0.17–0.66 mm RC per month, slightly lower than those for P. polymerus juveniles of 0.2–1.0mm
RC per month (Lewis & Chia 1981).
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For adult P. pollicipes growth rates were higher (3mm RC per year) compared with 1 or 2mm RC per year for P. polymerus (Lewis & Chia 1981).
Size at maturity, number of broods and number of eggs

Seasonal changes in the maturity of gonads of P. mitella (now Capitulum mitella) were observed by Yoshino & Konno (1987); they measured the diameter of the
oocytes throughout the year. They found no ovary in animals <13–15mm RC and no testes in those <9–11 mm RC. The animal was protandric. Batham (1945) found
that Pollicipes spinosus (now Calantica spinosa) had a breeding season in New Zealand from December to July with a maximum in February to April and she
estimated that it took about five years for adults to reach maturity. Although these cirripedes are no longer in the genus Pollicipes much of their behaviour is similar.
Pollicipes polymerus grows relatively slowly after the first year or so, depending on environmental conditions and reaches maturity in about five years. The animal
may be no more than half (15mm RC) its final size at this time. Barnes & Reese (1960) estimated that a full size adult may be 20–30yr old (see also Newman &
Abbott 1980). Examination of a large number of P. polymerus has shown that it is unusual for animals below about 14mm RC to contain egg lamellae. Bernard
(1988), on the west coast of Vancouver Island, found it was mature in its first year at 15mm RC but did not breed until the second year. At Monterey Bay, Hilgard
(1960) did not find egg lamellae in animals <17.2mm RC. Lewis & Chia (1981) also put the limit at about 17mm RC and breeding in the first year after settlement
(Table 16). On the southwestern coast of Portugal, Cardoso & Yule (1995) found egg lamellae in P. pollicipes of 8mm CH.
Table 16 P. polymerus: various details of reproductive period. Sizes, mm RC.
Place Size at breeding Breeding season Reference

Amphitrite Point, Vancouver Island, 49°N 15, breeds in 2nd year – Bernard 1988
San Juan Island, 48°30′N Late Apr. to early Oct. peak Jul. Lewis & Chia 1981
Edward’s Reef Eagle Point 17, majority at 32.5 14 to 27.5 (3 yr to breeding size) Begins at 8°C and rising. Stops at 9–10°C and falling
Bodega Head, 38°19′N – Peak Aug. Lewis & Chia 1981
San Francisco, 37°30′N – Late Mar. to early Jan. Lewis & Chia 1981
Monterey Bay, 36°40′N >17 Apr. to Dec.; peaks Jun., Sep., Dec.; at 12.3–17°C Hilgard 1960
Santa Barbara, 34°31′N (oil polluted) – All year, peak Feb. Straughan 1971
Goleta Point, 34°25′N >50% with eggs, 16 smallest 12, after 175 days – Page 1986
“Holly” oil platform 34°25′N >50% with eggs, 16 smallest 16, after 100 days – Page 1986
Santa Catalina Island 33°12′N 12, settlement to maturity 13 months – Cimberg 1978, 1981
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P. polymerus and P. pollicipes both appear to have a low temperature bar to reproduction. Hilgard (1960) has shown that the former species did not show
reproductive activity during four winter months when the mean shore, presumably air, temperature was below about 12°C. During the rest of the year when the
temperature reached 17°C a proportion of the animals had developing egg lamellae. At the peak of the breeding season young ova were always present in the ovary at
the same time as early stage embryos were in the mantle cavity. Even so, at the height of the breeding season she never found more than 60% of the mature animals
carrying egg lamellae. She, therefore, assumed that the reproductive period was determined by the time required for the development of the ova, which was longer than
that required to incubate the embryos. With the approach of winter small ova were present in the ovary with late stage lamellae in the mantle cavity—indicating the
beginning of the end of the reproductive period. In April and May the beginning of the reproductive period was signalled by the development of ova while no egg
lamellae were found in the mantle cavity. Cardoso & Yule (1995) found virtually no P. pollicipes carrying egg lamella during the winter months of October to March
on the southwestern coast of Portugal. From April to September from 20% to a maximum of 70% of the animals contained egg lamellae with a peak in July. Because of
local environmental conditions this maximum of 70% may be reduced to 40%, as in July 1993. On the northwestern coast of Spain, Molares et al. (1994b) found that
P. pollicipes had an annual gametogenic cycle with the main breeding season from March to September. The first peak was in March but the main one was in July to
September. These results are similar to those quoted by Barnes (1989, 1992; see below).
On the Pacific coast of north America the time and length of the breeding season of P. polymerus varies throughout the range of its distribution (Cimberg 1978,
1981, Lewis &
Figure 16 Pollicipes polymerus: percentage of animals with egg lamellae as a function of size of adult, = Goleta Point; ● = Holly.
(After Page 1986).
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Chia 1981). Page (1986) found differences in size and age at maturity depending on habitat. In particular he considered a rocky intertidal site at Goleta Point (34°25′
N) and a site 3km offshore on an oil and natural gas platform “Holly”. Factors influencing the growth at these two sites have been discussed on p. 358. He found that
the percentage of animals carrying egg lamellae increased with increase in size at both sites (Fig. 16). At Goleta Point the size for >50% to contain egg lamellae was
18mm capitulum height (CH) compared with 28mm CH at Holly. The smallest containing egg lamellae was 12mm CH at Goleta Point and 16mm CH at Holly. When
considering their growth rates this means that the animals are capable of carrying egg lamellae after 175 days at Goleta Point and after 100 days at Holly. Obviously the
formation of the gametes must have been initiated some time before this. There are three factors why the barnacles have a greater lifetime fecundity at Holly than at
Goleta Point. The adults attain larger sizes at Holly; they therefore produce larger egg lamellae, produce egg lamellae earlier in life, and they live longer and so produce
eggs longer.
Lewis & Chia (1981) also observed at San Juan Island (48°30′N) that higher barnacle growth rates were associated with a larger size of animals carrying egg
lamellae. The age of the onset of reproduction of 100 days at Holly and 175 days at Goleta Point (Page 1986), is nearer that given by Lewis & Chia (1981) of three
years than the estimated five years at Point Dune in southern California (Barnes & Reese 1959).
According to Cimberg (1978, 1981) there are two physiological races of P. polymerus on the Pacific coast of north America and these should be considered when
discussing the breeding seasons. The northern race shows maximum breeding activity at seawater temperatures of 14°C or less and the southern race at 20°C. He
distinguished three types of adults (Fig. 17).
Type 1 north of Point Conception, breeds in summer at seawater temperatures approaching 14 °C.
Type 2 at Point Latigo, south of Point Cenception, breeds in summer as seawater temperature increases up to 20°C.
Type 3 also found south of Point Conception—at Goleta Point and Santa Catalina Island breeds in winter as seawater temperature falls to about 14°C.
Figure 17 Point Conception: map of region showing nearshore currents during oceanic and upwelling seasons in relation to three sites
representing Types 1, 2 and 3 reproductive patterns. (After Cimberg 1981).
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Types 1 and 3 therefore breed at the same temperature but not in the same season and they constitute the northern race (“coldwater brooders”). Type 2 represents
the southern race (“warmwater brooders”). Types 1 and 3 may be similar because the southflowing California Current during oceanic and upwelling seasons (Fig. 17)
transports larvae from north of point Conception to the south. The season of maximum reproductive activity can change with the time of year when the temperature is
optimal for breeding. At Santa Catalina Island and Goleta Point the breeding seasons are six months apart from those of the population north of Point Conception, but
all are as the temperature approaches the same level. The P. polymerus population further south in Baja California breeds at about 20°C during the winter, that is six
months out of step with, but at the same temperature as the southern race at Latigo Point.
Point Conception is regarded as a biogeographical boundary for many taxa. Newman (1979b) noted that P. polymerus was the only species of 23 nearshore
temperate barnacles whose northern and southern limit did not end near Point Conception. There is, however, an interruption. At certain sites in the transition zone
between the two regions the distribution of these races of P. polymerus corresponds better to current patterns, which affect larval dispersal, than to temperature, which
affects the physiology of the adults.
The reproductive output of a species depends on how its survival and growth interact with reproduction. Breeding patterns of cirripedes and number of eggs
produced can vary with position within the intertidal zone (Barnes & Barnes 1968, Cimberg 1978, 1981, Lewis & Chia 1981). Page (1984) concentrated on an
examination of these two variables in P. polymerus collected from two localities at Goleta Point—an upper (+3 ft) and a lower (+0.5 ft) level—and from the intake
pipe of the seawater system (sws) at the Marine Station of the University of California at Santa Barbara. The first two sites were intertidal with varying degrees of wave
shock and exposure to air, but at the third (sws) site the animals were continuously submerged and the only turbulence was that caused by the intake water through a
pipe of 18cm diameter. The seawater temperature in the field ranged from a minimum of 13–14°C in December to May to a maximum of 20°C in August and
September. The temperature in the sws site was 3°C to 4°C cooler than the field seawater temperature in June to September. Page (1984) used animals with a CH of
> 15mm and made collections approximately monthly for 16 months (May to August of the following year). His results are summarized in Table 17. He also examined
the effect of time of submergence and found that the percentage of P. polymerus with egg lamellae was significantly greater after six weeks when continuously
submerged than when exposed to air for 9h in each 24h period. There was, however, no significant difference after only three weeks submergence.
The results of Page’s (1984) experiments emphasize the care that should be taken in comparative studies of reproductive patterns in intertidal cirripedes. His results
confirm the high variability in the percentage of animals with egg lamellae, size of egg lamellae, and estimates of reproductive effort that exist for P. polymerus.
Submergence time is always associated with more egg masses confirming the importance of feeding already noted in other localities (Cimberg 1978, 1981, Lewis &
Chia 1981). The temporal variation of animals carrying egg masses has also been related to water temperature (Hilgard 1960, Cimberg 1978, 1981, Lewis & Chia
1981). The highest percentages with egg masses at the intertidal sites of Page (1984) associated with lower water temperatures, and the high percentage in sws in
summer and autumn when water temperature was lower than the field temperature, together with data given by Page (1983), confirm that water temperatures of
approximately 14°C enhance reproduction of P. polymerus at Santa Barbara. Page’s (1984) comparisons of weights of egg lamellae were made on “standard”
animals of 16mm CH. Variability in
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Table 17 P. polymerus: fraction of time submerged per month (hours submerged/total hours) weightspecific investment per brood (slope of brood wt
body wt regression line), estimated annual number of broods produced per individual, reproductive effort (% body wt per yr as egg tissue), %
animals with egg lamellae. (After Page 1984)
+3 ft +0.5 ft sws
Jan.Jun. Jul.Dec.
Fraction of time submerged per month 0.30 0.83 1.00
% with egg lamellae 10–15 during 9 months 50–70, late winter and spring 20–30, late summer and autumn 70 all year round
Weight specific investment per brood, % body weight 4 12 7 21
Estimated number of broods 1 4.4 2.2 11
% reproductive effort 4 68 231
Weight of egg lamellae produced by animal of 16mm CH, mg 15–18 25–35 35, April 11, September 40–60
weight of egg masses (not standardized for barnacle weight) was examined at San Juan Island by Lewis & Chia (1981). In contrast to Page’s (1984) results, Lewis &
Chia (1981) found that the weight of egg mass was independent of the size of barnacle; it is obvious that this statement must be incorrect.
In order to compare egg production of different genera and species of cirripedes Barnes & Barnes (1968) devised a measure (NV)—increase in number of eggs per
50μg oven dry weight of animal (N) x egg volume (V). For P. polymerus and P. pollicipes the results are shown in Table 18. Compared with all the other cirripedes,
mostly operculates, that Barnes & Barnes (1968) examined the value for P. polymerus (NV=29) is very low. This species was collected at San Juan Island where the
animals only grow in a few very specialized habitats (Barnes & Reese 1960) and where conditions are less favourable than in the normal habitat—the exposed open
coast of north America. The value of NV=59 for P. pollicipes is of the same order but the higher value is in accord with the fact that conditions at Cabo Silleiro on the
Atlantic coast of Spain, where these animals were collected, are nearly optimal for the species. The lower metabolic rate as indicated by the oxygen uptake of both
isolated bodies and eggs of this genus has already been recorded by Barnes & Barnes (1959a,b).
Lewis & Chia (1981) say that “approximately seven times as many eggs are produced per brood by P. cornucopia [=P. pollicipes] at Cabo Silleiro and Gibraltar
(Barnes & Barnes 1968) compared with P. polymerus at San Juan Is”. This is incorrect, Barnes & Barnes
Table 18 Pollicipes species: size of stage I nauplius and comparison of egg production (N) and volume (V). (After Barnes & Barnes 1965, 1966, 1968).
P. polymerus P. pollicipes
Stage I nauplius, length μm 173 245
width, μm 82 –
6
volume, in 10 ml (V) 0.61 1.41
Increase in egg number per 50 μg oven dry body weight (N) 47.1 42
Product N×V 29 59
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Table 19 Pollicipes species: summary of breeding seasons, sizes and number of young, and number of broods per year. St. I=Stage I nauplius, o.d.=oven
dry. (After Barnes 1989, 1992).
Species Place Breeding season Size of young Number of eggs per Number of broods Reference
(LxB μm) brood per year
P. pollicipes S. Spain and Gibraltar 63% with eggs in July egg 245 15400 per 5mg o.d. Darwin 1851, Barnes &
body wt. Barnes 1966, 1968
Biarritz, France March to end Oct. – – – Barnes 1992
– – egg 154×78 – – Krüger 1940
Northwestern Spain March to September – – 1 to 2 Molares et al. 1994b
Southwestern Portugal April to September – – 1 to 3 Cardosa & Yule 1995
P. San Juan Island – ova 105×80 144000 to 288 000 2 to 4 Lewis 1975b, Lewis
polymerus &Chia 1981
Bodega Bay – egg 250×130 – 3 Lewis 1975b
Monterey Bay April, onwards, peaks in June, ova 100–127 104000 to 240000 4 to 5 Hilgard 1960
Sept., Dec.
Santa Monica – St. I 173×82 – – Barnes & Barnes 1965
Santa Barbara – St. I 207×114 – 8 Lewis & Chia 1981
Santa Barbara in area All year, peak Feb. – 80–120 8 Straughan 1971
polluted by oil
(1968) gave a figure of 15400 eggs for a moderate size P. pollicipes, which is seven times less, not more, than found for P. polymerus at San Juan Island. P.
pollicipes in Spain and at Gibraltar contained egg lamellae (63%) in July and at Biarittz, France 22% of the population contained egg lamellae in April, 85–90% in July
and August, and 16% in October (Barnes 1992). It is now known that 1–3 broods may be produced per year on the northwestern coast of Spain (Molares et al.
1994b) and in southwestern Portugal (Cardosa & Yule 1995, see also Table 19).
P. polymerus use a high proportion of their ingested energy in early life to produce somatic tissue and they reach a large size and are relatively older than many
operculates before reproductive activity begins. At this stage the energy requirement of 6–8cal day−1 (Page 1983) to maintain the large size results in a low investment
in reproduction.
Spermatogenesis and spermatozoa

Spermatogenesis has been investigated a little more than oogenesis in cirripedes according to Anderson (1994). Spermatozoa formation from the spermatogonium
through spermocytes and spermatids is similar in all the species so far examined. The electron microscope has played a great part in elucidating some of the processes
concerned. Dudley (1973) used such
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techniques to examine the synaptonemal polycomplexes in spermatocytes of P. polymerus from San Juan Island in Washington State, while Azevedo & Corral (1982)
made ultrastructural studies of the whole process of spermatogenesis as well as of the spermatozoa of P. pollicipes from Portugal. Dudley (1973) found analysis of the
spermatogenetic stages difficult because spermatogenesis of all the cells in one testicular lobe of P. polymerus was neither synchronous nor metachronic. This is
because in this hermaphrodite mature sperm are reported in some members of the population at any time during the year (Hilgard 1960). Dudley, however, does give
measurements of components of the synaptonemal complexes in the spermatocytes at the pachytene and diplotene stages of meiosis as well as micrographs of the
polycomplexes.
It has been long known that the spermatozoa of some cirripedes are filiform (Munn & Barnes 1970a). P. pollicipes from the Atlantic coast of Portugal near Oporto
was used by Azevedo & Corral (1982). At the end of the meiotic stages the spermatids have a spherical nucleus of about 4.9μm diameter and a volume of about
61.5μm3. The chromatin is evenly distributed and a nucleolus is present. In early spermatids the nucleus begins to elongate near the centriole region and the flagellum
elongates in the opposite direction. The chromatin becomes slightly condensed in the inner nuclear envelope near the axoneme region. One nucleolus is present in the
apical nuclear region. The Golgi apparatus is in the basal region near the proaccessory droplet. During this phase the mitochondrion moves to the base of the nucleus.
In middle spermatids the nucleus continues to elongate and the chromatin condenses more completely near the axoneme. Two sheets of 4–9 longitudinal microtubules
each surround the lateral zone of the nucleus. The mitochondrion is at the base of the nucleus and accompanies nuclear elongation while flagellogenesis takes place. The
nucleus becomes significantly elongated and filiform with dense chromatin in the third spermatids. The nucleus begins in the apical zone of the acrosomal basis with a
cap form having a doubly infolded zone where the centriole is found. There is a redundant nuclear vesicle that has a few nuclear pores with a diameter of 500–600 A.
At the end of nuclear maturation the membrane of the microtubules disappears as does the vesicle. The accessory droplet reaches its maximum length of about 14.4μm
and the matrix becomes denser and narrower. The accessory droplet is no longer than that of some other cirripedes so far examined, for example 8–9 μm long in
Balanus balanus, 10μm long in Balanus perforatus, but 20μm long in Semibalanus balanoides (Munn & Barnes 1970a). The spermatozoon of Pollicipes
pollicipes is similar to that described for other cirripedes (see Munn & Barnes 1970a,b, and references therein). It has a total length of 77.6μm, has an apical
acrosome, a centriole region, a nucleus (31.4μm long) and an accessory droplet region, an intermediate piece and tail. The axoneme has the usual 9+2 microtubular
pattern (Azevedo & Corral 1982). The accessory droplet (14.4μm long) begins about 7.1 μm below the acrosome. The sperm tail is about 45.4μm long with an end
piece 4μm in length. Some corresponding measurements of spermatozoa of other cirripedes have been given by Munn & Barnes (1970a).
In operculates the testes are confined to the thorax and prosoma, but in the pedunculates extensions may be found within the basal segments of the cirri and,
particularly in Pollicipes species, in the filamentary appendages. Hilgard (1960) followed development of the male gonads in P. polymerus as testes throughout the
body lost their translucent appearance and become opaque and white with sperm. She found that the diameter of the seminal vesicles was a good index of the maturity
of the male reproductive organs.
In nonparasite cirripedes the penis arises from an unarticulated support between the sixth pair of cirri and lies ventral to the anus. The structure of this support, the
pedicel, has been described for many species, including Pollicipes species by Barnes & Klepal (1971). The
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pedicel may be conveniently divided into two parts: a basal part and a distal part set at an angle to the former and directed rostrally. The penis arises at the distal end of
this distal part making a variable angle according to the characteristics of the distal part of the pedicel. The pedicel has a thickened band of chitin that varies with size
and species. The division between the basal and distal parts is termed the girdle.
The complexity and importance of the pedicel is greatest in operculates in which the extended penis is sharply angled down between the cirral bases. Barnes &
Klepal (1971) and Anderson (1994) give a detailed description of such a pedicel. The skeleton of the pedicel provides for muscle attachment and movement of the
pedicel and penis. There are also striated muscles that control the swinging and rotation of the penis in operculates in a horizontal orientation relative to the animal. In
pedunculates the skeletomuscular apparatus of the pedicel is considerably modified because the performance of the penis in these species is slower than in operculates
and has an arching manner with the base of the penis only slightly angled relative to the end of the thorax. According to Barnes & Klepal (1971) Pollicipes species,
particularly P. pollicipes, have a poorly developed pedicel structure.
Darwin (1851, 1854) has described the morphology and diversity of the penis in cirripedes in general; Klepal et al. (1972) were the first to elucidate its functional
organization in operculates. Cannon (1947) considered the penis of Lithotrya and its behaviour during copulation. The pedicels of Lithotrya and Pollicipes are very
similar and it is probable that the penes are also comparable. According to Cannon the penis of Lithotrya has only a retractor muscle and can only be extended by
erection. Unlike the cirri it only has one blood vessel, so blood forced into the penis from the epineural sinus can only escape the same way. During copulation the
occlusor muscles (socalled by Cannon) must first contract; this closes the exit from the body but allows blood from the capitulum into the body through the scutal
valves. Contraction increases blood pressure giving an erection of both cirri and penis. As long as occlusor muscles are contracted the penis will remain extended.
During copulation, the pedunculate penis slowly arches itself in a vertical orientation as it searches for a functional female. As far as the writer of this review knows this
has never yet been seen in Pollicipes species, but Kaufmann (1965) has photographs of the behaviour in the pedunculate Scalpellum scalpellum.
The paired canals from the seminal vesicles unite into a single duct at the base of the penis (Hilgard 1960). Darwin (1851) describes the penis of Pollicipes
pollicipes and P. polymerus as purple, short, with fine spines in tufts. The penis he found in a P. pollicipes was “a mere pointed papilla” but the animal was only
5.5mm CH and was not, therefore, mature. In both species the penis is muscular, short, inflexible, and heavily pigmented. The penis lies over the body of the animal
with its tip near the base of the first cirri, that is over the atrium (Barnes 1992). During copulation sperm is ejaculated into the mantle cavity of the functional female.
Sperm has been frequently seen covering the atrium of the oviduct in the mantle cavity of Pollicipes species; there is so far no published evidence of how it gets there
(Hilgard 1960, Barnes 1992). Gruvel (1893b) suggested selffertilization in P. pollicipes because he never observed any lengthening of the penis. Selffertilization
seems, however, to have been overruled by Hilgard (1960) and Lewis & Chia (1981). It should not be overlooked that Pollicipes can manoeuvre their capitula by
means of the peduncle (Barnes & Reese 1960) and an extendable penis may not, therefore, be as vital to copulation as it is in operculates.
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Oogenesis
In cirripedes oogenesis is relatively simple. Not much is known about the development of the ovary (Walker 1992). The ovary in Pollicipes species is in the peduncle
and the ovarian wall is lined with germinal epithelium. Here the oogonia undergo the preliminary stages of meiosis and produce the primary oocytes. Previtellogenesis
and vitellogenesis then follow, without involvement of nurse cells, until the breakdown of the germinal vesicle and the metaphase of the first meiotic division is reached.
The oocyte remains in this state until oviposition (Anderson 1994). Paired oviducts pass apically along the rostral side of the peduncle and mantle cavity before entering
the prosoma. The terminal parts of the oviducts, which are open at the bases of the first pairs of cirri, are modified as oviducal glands that become functional during
oviposition and fertilization. Cirripede oocytes are variable in shape and are surrounded by a vitelline membrane. They contain lipid droplets and membranebound yolk
granules containing carbohydrates and proteins. The cytoplasm also contains smooth endoplasmic reticulum, ribosomes, mitochondria, and glycogen granules (Lewis
1977). She gives an ova size of 105 x 80μm (length x width) (Lewis 1975b). Hilgard (1960) found that ova are virtually about the same size and at the same stage of
development throughout the ovarian tissue. An exception to this is when a new batch of tiny, immature ova appear in the ovary before the previous batch of very much
larger ova have been extruded.
Chromosomes
Chromosome numbers have been investigated in several cirripedes (see Austin 1987). The expected number of 32 was found by Nishikawa (1961) in Pollicipes
mitella (=Capitulum mitella) but so far there are no records of counts being made in the three Pollicipes species being considered in this review. Dudley (1973)
records a 2n=26 or 32 for some lepadiform cirripedes but says that the number for P. polymerus has not yet been determined.
Fertilization
Copulation
To fertilize the ova, spermatozoa have first to be deposited in the functional female by the penis of the functional male. Although this has been observed in many
cirripedes, so far there is no record in the literature of it having been seen in Pollicipes species. Even so sperm blobs are found in the mantle cavity over the atrium of
the oviduct. At the same time, ova are stimulated to pass up the oviduct into the oviducal sac and this is pushed out of the atrium but still attached to the oviduct. This
can easily be seen by carefully removing the sperm blob. Spermatozoa pass through the wall of the oviducal sac and fertilization takes place in the sac. There have been
differing ideas on how the spermatozoa get through into the sac (Klepal et al. 1977, Walker 1977, 1980, Anderson 1994) but it is now accepted that as the sac
expands, due to pressure from the ova inside it, small pores appear where the broken off cytoplasmic extensions were, and the spermatozoa penetrate the sac through
these pores probably using their acrosome. Once a pore has been penetrated it may be used by other spermatozoa or
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remained blocked by the underlying egg case (Walker 1977). At this stage when the sac is fully extended the wall is approximately 1μm thick and there may be 80000
pores of approximately 6μm diameter (Walker 1980).
It has been found that secretion from the oviducal gland is a powerful inducer of mobility in the spermatozoa after they have been deposited in the mantle cavity.
Lewis (1975b) found that only ejaculated spermatozoa taken from the mantle cavity of P. polymerus were capable of fertilizing ova in vitro. Spermatozoa taken from
the mantle cavity at the time ova are being released are without accessory droplets.
During copulation an excess of spermatozoa is produced and remains in the mantle cavity of the functional female as a gelatinous mass of coagulated sperm.
Examination by transmission electron microscopy shows that in this there has been extensive degeneration. Membranes have in part disintegrated, the axoneme may
have separated from the nucleus and some of the microtubules lie free in a structureless background. This has been found in P. pollicipes (Barnes et al. 1977). These
authors also followed the events in the mantle cavity immediately after copulation in P. pollicipes. Soon after the egg lamellae become free from the oviduct they are
manoeuvred to the sides of the body of the animal. This has to happen while the lamellae are still soft and pliable and before the increase of proteolytic activity due to
the formation of cross links in the membrane protein (Barnes & Barnes 1977).
Embryo development
Nussbaum (1889) first mentioned polar bodies of P. polymerus and in his monograph (Nussbaum 1890) described the first cleavage, polar body formation and
gastrulation in the same species. Groom’s (1894) stages of embryo development in Balanus perforates and Lepas anatifera have been used in reference to
Pollicipes embryos by Barnes & Barnes (1959b) although the emphasis was not on a description of the events during development. It should be recorded that there
are several occasions when Groom (1894) did not agree with Nussbaum’s (1890) observations on the embryo development of P. polymerus. This development from
fertilization through embryo and nauplii development to settlement has been followed in vitro by Lewis (1975b). She used eggs that had been fertilized in vitro and in
vivo and cultured them in filtered sea water at ambient seawater temperature of 13–15°C.
Soon after the sperm entry the ova become rounded and the fertilization membrane is raised and within a short time the first polar body is formed at the animal pole.
At this stage the fertilization membrane becomes sticky and appears to be responsible for the eggs adhering to each other. At the same time the eggs become elongated
to give the typical ovoid egg shape. There then follow peristaltic constrictions (Lewis et al. 1973). This stage is accompanied by ooplasmic segregation of yolk platelets
to the vegetal pole. There could be a reorganization of other materials, as well as yolk, at this stage. The egg membrane is first observed at points of the constriction
rings. The membrane is ovoid and strong; the second polar body is formed inside this egg membrane. After fertilization the volume of the embryo shrinks before it starts
to increase gradually as development proceeds. The stages of development, which are similar to those in other hermaphroditic cirripedes with planktotrophic nauplii,
are given in Table 20 as well as the time to reach that stage and the dimensions of the embryo. For comparison the times of development given by Nussbaum (1890)
and Hilgard (1960) are also included. Embryonic development in different conditions has been investigated by Lewis (1975a). These conditions involved light, aeration,
use of antimicrobial drugs, and water movement. Control cultures were kept in filtered sea water in about
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13–16h light per day and no drugs. When growth rates of the control embryos at 13–15°C were compared with those from under different conditions, it was found
that growth rates were faster in darkness and with aeration. These are the conditions most closely simulating the adult mantle cavity. Smaller portions of egg lamellae
and addition of antimicrobial drugs also promoted faster growth.
The times from fertilization to hatching are given in Table 20 and these are not very different from each other considering the different temperatures and methods of
determining the brooding time. Lewis (1975a) at 13–15°C gives 25.4 days (range 20.3–30.1 days), Hilgard (1960) at 13°C gives 29–31 days, Nussbaum (1890)
gives 20 days—the temperature was probably 21–23°C as this is what he quotes for some of his observations made in June 1888. It is not clear why Lewis (1975a)
gives Nussbaum’s temperature as 7°C in her table 7. Bernard (1988) gives a longer time of 50–60 days under natural conditions at Amphitrite Point, Vancouver
Island. This may be due to a lower temperature as he gives gametogenesis beginning at 9°C.
Nussbaum’s (1890) observations of the outline of the stage I nauplius and its internal organs, but not the setation, appear to agree with those of later workers (Lewis
1975b). In the natural population, hatching of stage I nauplii occurs in the mantle cavity of the adults
Table 20 P. polymerus: developmental timetable with embryo sizes (μm, measured inside egg membrane. (From Lewis 1975a, b). Data from other
authors added for comparison.—indicates no data available.
Mean embryo size, Time when event occurs for majority of embryos observed
Lewis (1975b) Lewis (1975a,b) Hilgard (1960) Nussbaum (1890)
Stage of embryo development Length Width 13–15°C 13°C 21–23°C
Ova, sperm added 105 80 0
Eggs roundup, fertilization membrane 100 100 0.3h
Fertilization membrane becomes sticky 100 100 0.6h
First peristaltic constriction rings 140 60 1.5h
First polar body, 2 constriction rings 140 60 2.0h
3–5 peristaltic constriction rings, eggs elongated 140 85 2.2h – First seen (x)
Elevation of egg membrane begins 140 85 2.2h –
1–2 peristaltic constriction rings 140 85 4.8h –
1–2 peristaltic constriction rings, 2nd polar body 140 90 5.6h x+6h
Egg membrane elevated 140 93 8.3h –
First cleavage 140 93 29.0h First seen (x) x+24h
Second cleavage 140 95 30.0h x+1 day
Third cleavage 145 100 37.5h – –
Fouth cleavage 145 100 55.2h – –
Fifth cleavage (31 cells) 130 90 60.0h – –
Sixth cleavage 135 90 65.0h – –
Gastrulation by epiboly 100 80 3.1 days – x+4 days
Division of internal yolky cell 110 80 4.4 days – x+7 days
Formation of mesoblast cells 130 90 5.1 days – –
Segmentation 120 90 6.8 days – x+8 days
Naupliar appendage buds 160 100 9.0 days x+10 days x+10 days
Origin of labrum and gut 185 115 11.3 days – –
Appendages with setae, origin of coelom 185 115 12.6 days – –
Naupliar eye, pronephros, horn gland 200 130 13.3 days x+16 days x+16 days
Limb movement within egg membrane, body transparent except for gut 220 135 16.7 days – –
Hatching 250 130 25.4 days x+29−31 days x+20 days
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and the nauplii are expelled in the mantle cavity current and propelled at least 3cm from the adult. In vitro, hatching takes place over a few days—the embryos at the
edges of the lamellae, where there is easier access to oxygen, hatch first. Hatching stage I nauplii were collected and raised in vitro through the six stages to cyprids by
Lewis (1975a).
The activity of several pentose shunt and glycolytic enzymes in developing eggs of P. polymerus from California have been investigated by Eastman (1968). The
rates of activity of glucose6phosphate dehydrogenase (G6PDh), 6phosphogluconate dehydrogenase (PGADh), phosphoglucoisomerase (PGI), and fructose
disphosphate aldolase (aldolase) have been compared at four stages during development of the eggs. There was major activity between the blastula stage and the
development of naupliar rudiments of all the enzymes except PGI. The ratio of aldolase to G6PDh activity was lowest in the early stages and highest at hatching. The
conclusion was that the pentose shunt may be important in glucose degradation during embryogenesis.
Naupliar development
The nauplii of P. polymerus are smaller than those of P. mitella (=Capitulum mitella; Yasugi 1937) and Pollicipes spinosus (=Calantica spinosa; Batham 1946).
This is to be expected in the last species, which has lecithotrophic nauplii. Pollicipes polymerus nauplii are planktotrophic and after the moult to stage II nauplii, food is
needed for the development to proceed further. Lewis (1975a) has investigated various factors affecting larval development in this species. She used multiplefactor
experimental designs to evaluate the effects of food type and concentration on survival of nauplii and their development. The experiments were done at a temperature
ranging from 12°13°C in spring and 13°16°C in summer. Several species of algae were fed to the nauplii but only three of those tested promoted growth to stage V
or older. Growth was fastest when a combination of Prorocentrum micans and Platymonas sp. was used and slowest when Platymonas sp. was replaced by a
pennate diatom. In this latter case most larvae did not moult beyond stage III. It is suggested that interactions between species of algae may affect their nutritional value
for Pollicipes polymerus larvae.
Lewis (1975b) gives the sizes of the nauplius stages of P. polymerus that had a total development time of 42 days at 12°C. Molares et al. (1994a) reported that P.
pollicipes required at least 23 days to reach the cyprid stage in laboratory cultures at 20°C. After 28 days, 28.3% of the larvae were cyprids. They fed the larvae on
Isochrysis sp. Lewis (1975a) found that Pollicipes polymerus nauplii did not get further than stage IV when fed on Isochrysis galbana. The individual stages of
Pollicipes pollicipes appear to be slightly bigger than in P. polymerus in every dimension except in the cyprid (Table 21). Lewis (1975b), however, had only one
cyprid giving a length of 425 μm and width 232 μm, whereas Molares et al. (1994a) had four cyprids and the range of length was 390–420 μm and of width was 210–
250 μm. Krüger (1940) quotes the size of a P. polymerus cyprid as 600×300μm.
A detailed description of the nauplius stages of P. polymerus is given by Lewis (1975b). This includes the eyes and carapace as well as the labrum. All balanoid
nauplii appear to have trilobate labra whereas in P. polymerus nauplii they are unilobate. She pays particular attention to the caudal spines and spinules on the
abdominal processes. She gives the setation formulae for the antennule, antenna, and mandible. The description of the naupliar stages of P. pollicipes by Molares et al.
(1994a) is not so detailed. Outline drawings are given but no setation formulae.
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Table 21 Pollicipes species: measurements of nauplius and cyprid stages (μm ± standard deviation); data from Molares et al. (1994a) rounded to
nearest whole number.
P. polymerus (Lewis 1975b) P. pollicipes (Molares et al. 1994a)

Nauplius Length Width Carapace length Length Width
stage Mean ±SD Mean +SD Mean ±SD Mean ±SD Mean ±SD
I 207 12 114 24 – – 245 28 151 24
II 315 10 185 8 – – 371 17 218 8
III 350 14 222 10 – – 406 8 245 7
IV 392 18 270 14 271 14 438 24 285 10
V 465 23 326 14 326 19 486 19 322 3
VI 542 30 390 22 383 22 558 15 367 12
Cyprid 425 – 232 – – – 403 15 231 20
Lewis (1975b) makes an estimate of larval dispersion from Bodega Bay, California; knowing the current speeds in the area (0.1–0.5 knots) and the time of larval
development (42 days), P. polymerus larvae could disperse up to 116–580 miles.
Reviews and summaries of naupliar development in various pedunculates and operculates have been published by Moyse (1987), Walker (1992), and Anderson
(1994). Each draws attention to differences between planktotrophic and lecithotrophic larvae—the former having larger growth increments between moults compared
with the latter. The general mechanism of feeding in planktotrophic larvae is discussed by Walker (1992) and particularly in lepadomorphs by Moyse (1987). Anderson
(1994) gives the dimensions of stage I and VI nauplii and the cyprid as well as development times for a host of thoracican larvae. At the time of his writing only the
larval stages of P. polymerus had been described but now those of P. pollicipes could be added to the list (Molares et al. 1994a, Kugele & Yule in prep.). Those of
P. elegans have still not been described.
Oviducal sac
When fertilized ova (the eggs) are found in the mantle cavity as paired egg lamellae it is evident that each lamella is surrounded by a single membrane. The nature and
formation of this has been a subject of controversy starting with Darwin (1851, 1854). He thought that the swelling at the base of each first cirrus was an acoustic
organ. Krohn (1859) showed it to be a terminal enlargement of the oviduct and suggested that this sac received the ova and was thereby expanded and forced through
the oviducal opening to give the egg lamella in the mantle cavity. Hoek (1883) at first proposed an alternative, but later (Hoek 1884) came to agree with Krohn (1859).
Gruvel (1893, 1905) described two entities, an atrium and a sac. Much of the early history on the nature of the oviducal sac has been summarized by Walley (1965)
and Klepal & Barnes (1977). The work of Walley completely confirmed Krohn’s (1859) interpretation of events. After examining a number of operculate species,
including Semibalanus balanoides, in detail she concluded that the form and function of the oviducal gland was the same in them as in the pedunculate Pollicipes
polymerus examined by Nussbaum (1890). He followed the course of the oviduct and general morphology of the sac in P. polymerus. At the beginning of the
secretory phase Walley (1965) states that material is shed into the lumen of the gland but is not incorporated into the wall of the sac. At the same time each cell of the
columnar epithelium is drawn into a cytoplasmic thread around which a cone of secretion is formed. As this extends towards the lumen to give a wall (15–20μm
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thick in Semibalanus balanoides) the epidermal cell shrinks to a small cuboidal element. The surface of the sac takes on a pitted appearance and in sections striations
are seen running from the pits through the wall. After distension by the ova to a thickness of approximately 1µm scattered spots are seen on the surface and Walley
(1965) was uncertain whether there were pores running through the wall sac or not. Nussbaum (1890) considered that the striations, seen in sections to pass through
the membrane of the sac of Pollicipes polymerus, indicated the presence of pores. Even though in this genus the striations are particularly clear there have been
instances in which striations of similar appearance did not indicate pores. Whether pores are present or not is of considerable interest as regards the access of
spermatozoa to the ova. In the absence of pores the spermatozoa would have to penetrate the sac wall before fertilization was possible.
Barnes & Barnes (1977) found that the oviducal sacs of both P. polymerus and P. pollicipes could be easily obtained by gentle pressure on the bases of the first
cirri within which they lie at a time when the animals are in a breeding condition (see also Barnes 1992). Klepal & Barnes (1977) found that, as in Semibalanus
balanoides, the sac in Pollicipes is secreted around cytoplasmic filaments arising from the epithelial cells. These filaments are longer than in Semibalanus balanoides
and the thickness of the sac is consequently greater (approximately 40μm). Because of this and its apparent rigidity the sac of Pollicipes tends to preserve its shape
(Fig. 18). The wall is in part folded and basally, towards the entrance of the oviduct, it spreads to form a convoluted rim. In transverse sections of the sac striations
Figure 18 Pollicipes pollicipes: oviducal sac removed from end of oviduct of a mature animal immediately before copulation.
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are seen running through the wall secretion and terminating in “pores” at different levels. The same is seen in sections of the outer part of the wall. Klepal & Barnes
(1977) examined the oviducal sacs of Pollicipes in great detail by both scanning and transmission electron microscopy. They give numerous figures and dimensions of
all the appropriate components. These authors agree with Nussbaum (1890) that the fine lines seen in sections of the wall of P. polymerus sacs represent pores and
that the “spotted” appearance of the sac and striations seen by Walley (1965) do, as she suggested, correspond to the position of the cytoplasmic filaments.
During embryonic development the eggs containing the embryos are enclosed in the vastly stretched oviducal sac and are incubated in the mantle cavity of the adult.
In this way the developing eggs are prevented from being carried out of the mantle cavity during respiration or feeding activities. At about the same time as the embryos
mature the sac begins to disintegrate. Barnes & Barnes (1977) investigated three possible causes of this breakdown. The first, bacterial activity within the mantle cavity
was disproved as a primary cause. The second, namely a substance released by the adult, is equally unlikely because egg lamellae separated from the adults have been
cultured through to the liberation of nauplii and in these the sac also breaks down. The simplest explanation is the third, that at the time the embryos are ready to hatch
they produce a material, presumably enzymatic in nature, which acts on the sac and destroys it. Such a substance does not, however, affect the egg cases and therefore
should not be confused with the socalled hatching substance.
The oviducal sac is proteinaceous (see below, Barnes & Blackstock 1977b) and protease (pronase) has often been used to free eggs from egg lamellae when egg
counts were needed. The effect of protease on egg lamellae at different stages of development in several species, including P. pollicipes has been examined by Barnes
& Barnes (1977). The time taken to breakdown the sac in P. pollicipes increased from about 2h initially to 7–8h when the egg lamellae were rigid and the eggs many
celled. Later the time decreased until it took <1h when hatching of the embryos was imminent. These times are longer than for the other species and may only indicate
differences in the protein composition of the sacs (see Barnes & Blackstock 1977b). The times are, however, a function of the state of embryonic development rather
than “real” time. The fact that the sac is almost pure protein together with the observed changes in rigidity and change in resistance to protease of the egg lamellae
indicates that polymerization must take place and probably starts in the oviducal gland (Barnes & Blackstock 1977b). The breakdown of the sac by protease indicates
removal of the sac by a proteolytic enzyme produced by the embryos themselves. In order to avoid confusion with the hatching substance Barnes & Barnes (1977)
gave the name “ovisacase” to the cirripede enzyme responsible for proteolysis of the oviducal sac.
The properties of ovisacase have been examined by Barnes & Blackstock (1977a). It was found to be relatively heat stable and may be stored at low temperatures
for long periods without loss of activity. It differs from other known hatching enzymes in having optimal activity in the acid range (approximately pH 3.0). It has a wide
range of proteolytic activity and there is little need for metallic activation. Although showing a wide range of activity to synthetic peptides, as regards activity towards the
sac material there may be considerable enzyme specificity in cirripedes.
The aminoacid component of the protein (Barnes & Blackstock 1977b) in the oviducal sac of Pollicipes pollicipes is different from that of Semibalanus
balanoides (Table 22). Glutamic acid, alanine, and leucine, the major components in Pollicipes, make up 40.3% of the total; glycine, alanine, glutamic acid and serine
equal 47.7% in Semibalanus balanoides. The most striking differences are higher glutamic acid and lower glycine in Pollicipes. These
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Table 22 P. pollicipes: aminoacid composition of oviducal sacs compared with those found in Semibalanus balanoides. Tr=trace. (After Barnes &
Blackstock 1977b).
Amino acid, residues

100–1 residues P. pollicipes S. balanoides
Aspartic acid 5.8 7.9
Threonine 5.2 5.9
Serine 3.9 9.4
Glutamic acid 14.3 9.6
Proline 6.8 5.3
Glycine 7.3 18.7
Alanine 11.0 10.0
1/2 Cystine Tr Tr
Valine 9.1 6.5
Methionine 1.8 –
Isoleucine 3.4 3.8
Leucine 15.0 5.8
Tyrosine 1.0 2.1
Ph. alanine 2.6 2.9
Lysine 1.9 3.8
Histidine 1.8 2.2
Arginine 9.0 6.1
differences may be related to species specificity of the two ovisacases (Barnes & Blackstock 1977a) and to less complete breakdown of the oviducal sac in Pollicipes
in natural and experimental conditions (Barnes & Barnes, 1977).
As there is no evidence of any pores in the egg case (Barnes et al. 1977) in what form does the enzyme pass through the egg case? How is the enzyme activated? As
both egg case and oviducal sac are largely protein, how is the former able to withstand proteolysis? Substrate specificity may be the answer to this.
Ultrastructural investigations
The formation of the three envelopes that cover the cirripede embryo have already been mentioned—the oviducal sac that forms the outer covering of the whole egg
lamella, the fertilization membrane, and the egg case. After entry of the spermatozoon into the ova fertilization takes place and the fertilization membrane is raised.
Electrondense granular material from the vesicular bodies in the cortex is incorporated into the vitelline layer as it is transformed into the fertilization membrane. This
has been followed in P. pollicipes by Klepal et al. (1979) using transmission electron microcopy. They give figures and measurements of the various stages of
development. The separation of the fertilization membrane from the surface of the egg occurs at about the time of the first meiotic division. The oocytes with their
vitelline layer are not adherent to one another when freed from the ovary; this is essential for their further liberation and movement through the oviduct into the oviducal
sac. The adherence of the eggs after fertilization is due to the “sticky” nature of the fertilization membrane and it would seem, therefore, that this may be, in part,
because of cortical material incorporated into the vitelline layer as the fertilization membrane is produced. Lewis (1977) also examined the formation of the fertilization
membrane in P. polymerus. The first polar
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body becomes separated from the surface of the egg within the fertilization membrane. This membrane then becomes organized into two layers and these are concerned
in the early formation of the egg case.
A series of contractions begins after the first polar body is given off and at the same time an envelope is formed round the developing egg. The peristaltic
constrictions exhibited by cirripede embryos during early development, and noticed by Groom (1894) in P. polymerus, have intrigued Lewis et al. (1973) and Lewis
(1977). Groom (1894) said the waves of constriction were shown before the protoplasm and yolk had separated. These constrictions slowly moved as rings from the
animal to the vegetal pole of the newly fertilized egg. Lewis et al. (1973) used timelapse cinematography to follow these movements and attempted to find some
evidence of their causal mechanism by the use of different chemical inhibitors. They showed that it took about 8.5min for a constriction ring to travel about 85% of the
egg’s length (total length 140μm). The rings also travelled faster in the animal half than in the vegetal half; the velocity also decreased with age of embryo. The rates of
peristaltic motion were independent of the number of rings present; the interval between rings reaching the vegetal pole was 3min in eggs having 3 or 4 rings. The
ultrastructure of the egg of P. polymerus with peristaltic constrictions was investigated by Lewis (1977). In her paper she describes the fine structure of the egg during
the formation of the first polar body and peristalsis.
Dense bodies were produced by the Golgi during the formation of the polar body and extracellular plaques were observed. These were thin microfilaments (40–60
A) in the egg adjacent to the polar body. Extracellular spheres, flocculent material, and filaments were seen in eggs with peristaltic movement. At the level of the
constrictions intracellularly there were numbers of multivesiculate bodies, glycogen granules, mitchondria, proteincarbonate, and lipid yolk bodies. Microfilaments in the
cortical area were found exclusively in the peristaltic constriction rings. Filaments were orientated primarily in the meshwork, and some circumferentially placed ones
were found in rings near the vegetal pole. The space between the constriction and the elevated egg membrane contains extended microvilli. Possible functions of the
peristalsis in cirripede eggs have been suggested by Lewis et al. (1973) and Lewis (1977). Elevation of the egg membrane may be related to the constrictions but unless
secretory activity occurs as a result of constriction there is not likely to be a direct connection. Egg elongation is certainly accomplished during peristaltic contractions.
Another possible function may be the predetermination of the first cleavage furrow. Lewis (1977) describes a model to explain peristalsis in P. polymerus.
When the contractions cease the second polar body is given off within the envelope. Occasionally it is possible to find eggs with two polar bodies, one inside and the
other outside this envelope. This consists of an outer more electrondense layer (0.02–0.03 μm thick) and a second layer (0.06–0.07 μm thick). This decreases in
electron density as it is laid down and eventually becomes a finely granular layer (0.16–0.18μm thick). When the embryo reaches about the limb bud stage a second
envelope is formed round the eggs, termed the embryonic sheath by Klepal et al (1979). This also eventually consists of two layers: an outer electrondense layer
(approximately 0.01 μm thick) and an inner finely granular layer (approximately 0.22–0.26μm thick). These two layers are similar in appearance to the two layers of
the first envelope. The two envelopes, now approximately 0.5μm thick, form the egg case. By hatching time it will have become 0.3μm thick due to stretching by the
developing embryo. As hatching approaches, the inner layer of the embryonic sheath begins to disintegrate and eventually disappears. The stage I naplius emerges
eventually from the anterior (blunt) end of the egg case and there is evidence that just before hatching the embryonic sheath separates from
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the outer sheath (first envelope of the embryo) and expands bursting the outer sheath; the nauplius has then only to push through the stretched remains of the embryonic
sheath.
The movement of the ripe embryo before hatching by flexing the caudal region must have an abrasive action on the outer layer of the embryonic sheath. Barnes &
Barnes (1977) have shown that an enzyme, ovisacase, is secreted by the embryo and this attacks the oviducal sac surrounding the egg lamellae. This enzyme may be
produced at the same time as the embryonic sheath. Klepal et al. (1979) speculated that coated vesicles seen inside the developing egg case, and later outside it, may
contain the ovisacase because it passes from the inside to the outside of the egg case without having any effect on it. It seems that another enzymatic process must be
responsible for the breakdown of the inner layer of the embryonic sheath. In their analysis of egg cases, which were probably contaminated with remnants of the inner
layer of the embryonic sheath, Barnes & Blackstock (1977b) found an unexpected quantity of iron. This was not contamination and they suggested that an ironbound
protein may protect an egg case from enzymatic attack in contrast to that caused in the embryonic sheath and the oviducal sac.
It is common knowledge that a socalled hatching substance can enhance hatching of cirripede eggs. This substance is contained in homogenates of adult cirripede
tissue. Recent work has shown that the action of such homogenates is due to two groups or types of eicosanoids (Song et al. 1990). These authors extracted
eicosanoids from homogenates of several cirripede species including P. pollicipes from Portugal. All the extracts contained a monohydroxy fatty acid fraction (MHFA)
and a trihydroxy fatty acid fraction (THFA). The results of the bioassay of MHFA and THFA fractions on the egg masses of four species of operculates showed the
presence of hatching activity in P. pollicipes extracts that had not been previously recorded.
Endolithic algae
The occurrence of the Conchocelisphase of a species of Porphyra in living Pollicipes pollicipes from Roscoff has been recorded by Drew & Richards (1953). At
that time this was the first record of the occurrence of this phase of Porphyra on the coast of France and it was found in calcareous parts of an attached living organism
found in a specific type of habitat where wave action is strong. This was of ecological interest in consideration of the life history of the Porphyra species to which this
Conchocelisphase belonged.
The infection of Pollicipes pollicipes seems to have been almost entirely restricted to the peduncular scales with only limited infections on the scutum and a lateral
plate of one animal. The barnacles examined were sexually mature. There was dense growth of the Conchocelisphase suggesting that the infection was not new.
Infection was only found in plates and peduncular scales where the protective cuticular layer had been destroyed.
Other algae were found on the outside of the scales and at least one other unidentified endolithic alga was present in the scales. Algae, bryozoans and lamellibranchs
have frequently been found attached to the peduncles of large Pollicipes pollicipes (Broch 1927).
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Predation
Distribution and abundance of intertidal organisms can be influenced by several factors of which predation is one. Sometimes this is not considered as seriously as it
should be. The predators may be other intertidal organisms, fish, birds or humans. The effect of collection by humans for food and the possibility of organizing fisheries
for Pollicipes on a commercial basis are considered on pp. 381–3.
Bernard (1988) recorded predation of P. polymerus on a rocky promontory on Vancouver Island, British Columbia. After a settlement of cyprids numerous young
adults were found on peduncles of older adults, algal fronds, and coralline algae in tide pools as well as on cleared sites. After six weeks almost all the young barnacles
had disappeared due to predation by small crabs and polychaetes. In similar studies he ranged the potential predators in order of significance as Pisaster ochraceous,
muricid snails Nucella emarginata and N. lamellosa, and small pagurid crabs. Palmer (1984) also found N. emarginata preying on Pollicipes polymerus. According
to West (1986) the barnacle is drilled between its rostral and scutal plates by the predator. Hui (1983a) thought predation by crabs might be responsible for
contraction of the peduncle in Pollicipes pollicipes. Bernard (1988) suggests that predation may limit Pollicipes species to the upper third of the tidal zone where tidal
surges are strong. Hoffman (1989) often found the nemertean Emplectonema gracile, foraging on young animals; the worms were wrapped round small animals many
of which often lacked capitula. He also found large numbers of polyclad flatworms, crabs, amphipods, and opisthobranch molluscs in the dense clumps of Pollicipes
polymerus. Any of these organisms were possible predators. He thought that the lower midintertidal area was best for survival. Some balance has presumably to be
struck by local conditions as to which part of the intertidal guarantees the best recruitment.
As has already been pointed out Pollicipes species are often associated with mussels, and mussels are frequently subjected to predation by seabirds. Marsh (1986)
found Pollicipes in pellets regurgitated by gulls. Presumably this was P. polymerus as the work was done on the coast of Oregon, USA. During autumn and winter
birds in flocks of up to 100 are common on the west coast of north America. Mussels and Pollicipes form a major part of the winter diet of surfbirds, Aphriza
virgata, and gulls, principally Larus occidentalis and L. glaucescens. From November to March, surfbirds consume the smaller Pollicipes polymerus and mussels.
The surfbirds are generally absent for the rest of the year. The gulls consume mostly the larger Pollicipes in winter; at other times they primarily feed offshore. In spite
of these, and other incidental observations, Meese (1993) made the first detailed and experimental investigation of the predation effect of birds on the distribution and
abundance of P. polymerus. His experimental area was the outer coast of the Bodega Marine Reserve in California. This rocky intertidal zone is composed of sloping
granite and is dissected by surge channels and tide pools. Meese used cages to exclude avian predators. The cages were raised about 3–4cm above the ground thus
allowing access to intertidal predators such as Pisaster ochraceous and Nucella species. The mesh used in the cages was too coarse to exclude the birds’ beaks but
their heads could not fit inside the mesh. This allowed foraging inside the perimeter of the exclusion plots. There may have been the occasional forage by intertidal fish
but usually sea conditions were so rough as to preclude this. The cages were deployed from April through to August of the following year. For the first fourteen months
they were examined monthly and thereafter every two to three months for 21 months. Twice monthly bird surveys were made with binoculars during periods of low
tides for almost four years.
The Pollicipes usually formed 30–50% of the total plot cover and tended to decrease with
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time. Control areas showed significant reductions in Pollicipes cover during the winter. There was, however, significantly more lost from the control than the exclusion
plots. This suggests that predation by birds had a relatively large impact during the winter. At the Bodega site Pollicipes is not restricted to the narrow range of habitats
with specific hydrodynamical features, and in the time of experiment the species was not outcompeted by Mytilus californianus. Although morphological and physical
factors (Barnes & Reese 1960) and interspecific competition for space (Paine 1974) may affect distribution and abundance patterns of Pollicipes in some places on
the Pacific coast of north America, in northern California predation by birds must also be considered.
Barnes & Reese (1960) described the aggregative “rosette” pattern of P. polymerus (see Fig. 1 p. 306) and suggested that this provided protection from predation
but did not say which predators. Meese (1993) speculated that a tight rosette formation was the result of the “shadow reflex” (Gwilliam 1963) initiated by the shading
of gulls. This produced more tightly packed rosettes than ones in which the peduncles were elongated. Many tight rosettes may be attacked before a foraging gull can
remove an individual barnacle, but once this happens the whole rosette is susceptible to further predation by that and other gulls. Whole rosettes can be consumed in
this way thus leaving clear spaces and causing patchiness in the distribution pattern of P. polymerus.
Pollution
The Santa Barbara oil spill of 1969 was caused by a massive flow of oil from an offshore drilling accident. Crude oil has continued to seep from the sea floor even
though the initial outflow has been stopped; this seepage is likely to continue. There is oil and gas activity at a small natural seep at 60ft depth. This knowledge and the
anticipation of increased domestic and industrial pollution along the Californian coast has prompted systematic observations of the effect on marine fauna and flora from
Santa Barbara to San Diego (Foster et al. 1971).
Preliminary reports have been presented by Straughan (1969a,b) on the breeding activities of several intertidal species including P. polymerus. At the time of the
first collections no reproductive activity was reported in this species. Straughan (1971) made a more detailed survey to determine the effects of sublethal doses of oil
from natural seepage and of oil from the single spill. Sedentary species such as P. polymerus were unable to escape the pollution. The abundance of this species was
visually assessed; it was less abundant on stable rocks south of Santa Barbara Channel than at nonseep areas within and to the north of the Channel. In this region P.
polymerus breeds throughout the year but this breeding was reduced in polluted areas. Breeding in two balanoid barnacles did not seem to be so affected but the
stalked barnacles occurred here in the lower intertidal areas and so were exposed longer to the effects of oil. In midintertidal zones oil tended to stick to the capitular
plates of P. polymerus and they were killed when the oil became thick enough to smother them. In many cases the cirri were covered with dried oil. In areas where
there was sufficient exposure to wave action the oil might be washed off before it had a chance to dry (Foster et al. 1971). P. polymerus settlement was found on un
oiled, but never on moderately or heavily oiled areas of peduncles. Liquid oil tended to accumulate in the clumps of Pollicipes and this could mask any conspecific
chemical attractant making the surface unattractive to cyprids.
So far populations of Pollicipes have survived in the presence of moderate oil pollution because of planktonic renewal from nearby unpolluted sites. At some stage,
however, plank
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tonic renewal will be affected and the reduction in breeding and space for settlement will reach the level where the population cannot be maintained.
Nuclear power reactors, nuclear fuel reprocessing, and the testing of nuclear weapons release tritium in liquid effluent or into atmosphere. This has resulted in
increases over that present in the world as a result of natural production. The biological impact of such increases has been investigated mostly on mammals. Studies of
the effects on invertebrates have been less rigorous; deleterious effects have been found on oysters and Artemia nauplii. Abbott & Mix (1979) have investigated the
effect on the development of Pollicipes polymerus nauplii. They used a “moulting index” [(number of post stage I nauplii)/(number of stage I nauplii + number of post
stage I nauplii) 100]. Apart from the anomalous result in the control experiment, they found a great reduction in the moulting index as concentration of tritiated water
increased. They concluded that as harmful effects were caused at about 1.5×10−5μCiml−1 there should be a change in the permissible maximum concentration of
3×10−3μCiml−1 in water for the general population.
Commercial exploitation
The soft parts of several species of cirripedes have been a source of food for local inhabitants for generations. It is usually the ovary or eggs that are cooked and eaten
but in some places, where very large genera are available, the muscles of the prosoma may also be eaten. The ovary of all three species of Pollicipes is much sought
after in areas where they are common.
P. elegans
There is a small localized fishery for P. elegans in Costa Rica but apart from this little is known about the consumption of this tropical Pacific species (Bernard 1988).
P. polymerus
Native people living in the coastal areas of North America used P. polymerus as food but more sparingly than P. pollicipes was, and is, used in Europe. If the
muscular stalk of P. polymerus is boiled or steamed for 20 minutes and the skin removed the remainder, red in colour, is said to be moderately good eating and tastes
like lobster (Cornwall 1955b, Bernard 1988). Gibbons (1964, cited in Newman & Abbott 1980) even gives recipes.
The European P. pollicipes is now in short supply (see below) so there is a demand for P. polymerus to be exported from North America to Spain and Portugal
(Newman & Abbott 1980). The demand in Spain is estimated at 2000 metric tonnes per year and there is a substantial demand in Portugal for animals with stalks more
than 4cm long. P. polymerus is the largest of the three Pollicipes species and the stalk can be up to 10cm long. It is abundant in places where it can be harvested and
it could form the basis of a fishery if storage and transport problems could be resolved. In British Columbia a sharpened metal bar is used for harvesting on friable
argillitic rock but a hammer and chisel are necessary on sandstone substrata. Test shipments have been made. Numerous scientific permits to harvest the barnacles
were issued in 1977 by the Department of Fisheries and Oceans. In 1978 about 25 metric
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tonnes were harvested from southwestern Vancouver Island and some were flown alive to Spain. Quality control and transport problems led to the demise of this
venture but activity at a reduced level continued until 1984 when two attempts were again made to initiate a fishery; they also failed. Activity was renewed in 1985 and
1986 and in 1987 several companies in British Columbia successfully exported more than 20 metric tonnes by air to Europe (Bernard 1988).
The harvesting of material on this scale does, however, present problems for the local species such as what is the recovery rate of the cleared areas? Bernard (1988)
has made a study of a rocky promontory on the southwestern coast of Vancouver Island (see p. 358). The conclusions were not promising for a viable fishery and
export of live animals to Europe. There is the problem of separating the animals from the hard rocks found on the shores of British Columbia without damaging the soft
tissues. Even the lowest mortality and decomposition can ruin a whole shipment of live animals. Harvesting the barnacles from mussel shells would be easier but less
than 2% of the population in British Columbia are attached to such shells.
Survival of Pollicipes could be improved by wet storage but given the delays in air transport it seems that despatch frozen would be the only satisfactory way.
Crustacean flesh is only slightly affected by quick freezing and it has no obvious detrimental effect on the Pollicipes stalk (Bernard 1988 and references therein). It is
unlikely, however, that the European market will pay the prices for a frozen product. Although P. polymerus is abundant in British Columbia, many places are exposed
to storms or of difficult access and so it is difficult to estimate the standing stock that could be harvested. There is also evidence that harvested areas are not readily re
settled and so the sustainable yield for any fishery is expected to be a very small fraction of the standing stock. The inaccessibility of most of the harvestable stock gives
the species adequate protection for the future (Bernard 1988).
P. pollicipes
P. pollicipes has been known for over two centuries by Algerian fishermen who gave the barnacle the imaginative name of “patte de chèvre” or “pied de chèvre”—pad
or foot of a goat (Llabador 1937). It has, however, been known and used since prehistoric times. Its calcified plates have been found during excavation of grottoes and
caves. Doumergue (1921) found them in Neolithic hearths in caves near Oran. Such finds caused some speculation as to their origin. Were the animals eaten and the
plates discarded or were they used as ornaments or jewellery? None of the plates found had been pierced and so Doumergue (1921) assumed that the animals had
been a source of food. Such remains have also been found in caves at Cap Spartel near Tangiers.
In many places in Europe P. pollicipes does not form dense masses occupying much of the midlittoral area of the coast as does P. polymerus on the Pacific coast
of North America. This scarcity may be due, in part, to long stretches of unsuitable habitat on European coasts or, perhaps more likely, to long continued overfishing
(FischerPiette & Prenant 1956). On these coasts this cirripede has been, and still is, much sought after for food. Newman & Abbott (1980) say that the barnacle is
cooked and served as a delicacy in Spain and Portugal. Darwin (1851) has a footnote about P. pollicipes being eaten as a food in Brittany. This is confirmed for
Brittany by GuerinGanivet (1907), Joubin (1907), and GuerinGanivet & Legendre (1909). There are vertical cliffs and reefs that are inaccessible except by boat
where fishermen make their collections (FischerPiette & Prenant 1957). The stalks of the
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barnacles (containing the ovary) are cooked, skinned and the pinkish ovary eaten. Joubin (1907) did not share the people’s enthusiasm for it and GuerinGanivet &
Legendre (1909) found it tough but with an agreeable flavour. The author of this review having also tasted it, is in total agreement—a lot of work and very little reward!
There is, however, a great desire in Spain, and in Portugal particularly, to encourage the cultivation of P. pollicipes and to increase commercial fisheries for it (Cruz
1993, Molares et al. 1994a). Consumption in Spain alone is estimated at 2000 metric tonnes per year, that is a current market value of about 63 million dollars
(Proverbs 1979 cited in Bernard 1988, Molares et al. 1994a). This interest has prompted a renewed effort to study the biology and ecology of this species with a view
to exploiting it commercially in Europe.
Stocks in Spain have been so depleted that there are now strict conservation measures. Since 1970 the demand has been met partly by importing the animals from
France and Morocco but these sources are now exhausted due to overfishing. Attention has therefore turned to the North American species (see above) but there is
some resistance by the Europeans because of the unfamiliar taste and tougher meat of the American species.
Conclusions
The three Pollicipes species have a similar appearance although a few details may differ. There is a capitulum, well endowed with calcified plates, that protects the soft
parts of the animal and contains the developing egg lamellae when present. The ovary is contained in a strong peduncle which has a thick integument again covered with
small calcified scales. When the peduncle is contracted the whole appearance is of a very sturdy animal.
The favoured habitats for this genus are areas of high exposure where the waves surge in channels or up gullies in steep cliffs and the backwash is strong. In Europe
P. pollicipes may be found in caves at the end of channels where the waves break with extreme violence. The animal needs the force of the incoming water to stimulate
the opercular valves to open. At the same time the peduncle orientates the capitulum so that the cirral net faces the backwash of water. Except in juveniles, there is no
cirral beating action as in many operculates. The cirri remain extended with only individual cirrus reacting to stimulation by food particles. This extension is due to
hydrostatic pressure. The structure of the peduncle is such that it forms what some workers call a hydrostatic pressure cylinder. The pressure and musculature of the
peduncle give it rigidity but it still remains flexible enough to reorientate the capitulum, if necessary, due to changes of water currents.
In an intertidal cirripede gaseous exchange has to take place in water and when exposed to air. In P. polymerus there is an increase in aerobic metabolism on
emersion in contrast to the reduced oxygen uptake in operculates under such conditions. This may be owing to the lack of rhythmical cirral beating in Pollicipes so that
cessation on exposure to air has little effect on the respiratory needs. For oxygen uptake to remain high on emersion it has been suggested that gaseous exchange
across the wall of the peduncle may be involved. This would require an efficient high pressure haemolymph circulation such as that found in Pollicipes. That aerobic
metabolism can be enhanced on exposure to air has been suggested as a probable contributor to the success of these pedunculates in the intertidal community.
The surface of the peduncle is also an asset in providing an additional area for evaporation during emersion to cool the animal during heat stress.
Communities of Pollicipes sp. and Mytilus sp. are frequently found on the Pacific coast of
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north America and in Europe and on the west coast of Africa, but neither organism seems to outcompete the other. In denuded areas Pollicipes may be the first to
establish itself but after several years Mytilus is also established. Where the terrain is very exposed and on steep or vertical surfaces Pollicipes seems at an advantage.
This may be because of stronger adherence to the substratum or the protection ensured within fissures in the rock. In such cases Mytilus remains at the lower levels.
Predation may be a problem in the very young stages but later the rapid closure of the opercular valves and contraction of the peduncle give good protection. The habit
of forming clusters of adults also gives added protection.
As has been seen many thousands of eggs are produced in several broods each year and the adult peduncles provide ideal surfaces for the settling cyprids. Once
again the habit of forming clusters of adults gives protection to the juveniles.
The commercial exploitation of Pollicipes, particularly in Europe, has encouraged a renewed interest in these pedunculate cirripedes and hopefully it will also
increase some fundamental research on their biology and ecology.
To be successful in the intertidal region a marine animal must be capable of withstanding the rigours of that environment. These are manifold; the ability to adjust to
periods of submersion and emersion, fluctuating temperatures of water and air, desiccation stress when exposed to air, changes in respiratory processes, and gaseous
exchange is essential. There are also physical hazards such as the pounding by waves in exposed situations, competition from other animals, effects of pollution, and
predators. In addition, an animal must be capable of reproducing itself successfully. The genus Pollicipes seems to be well adapted to meet all these conditions.
Acknowledgements
I am grateful to Professor W.Klepal for reading parts of the manuscript, to Dr A.B.Yule for allowing me access to work “in press” and “in preparation”, to Dr
A.D.Ansell for advice and help with the figures, to Miss E.Walton for assistance in obtaining obscure literature, and to Mrs S.D.Pearson for typing the manuscript.
I am glad that in writing this review I have been able to incorporate notes prepared by my husband, Harold Barnes, before his untimely death in 1978.1 also wish to
express my appreciation to colleagues and friends worldwide who have encouraged me in my cirripede work during the intervening years.
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COASTAL FISHERIES IN THE PACIFIC ISLANDS

P.DALZELL,1 T.J.H.ADAMS1 & N.V.C.POLUNIN2
1Resource Assessment Section, Coastal Fisheries Programme, South Pacific Commission BP D5, Noumea, New Caledonia.
2Centre for Tropical Coastal Management Studies, Department of Marine Sciences and Coastal Management, University of Newcastle upon
Tyne, England.

Abstract Coastal fisheries in the South Pacific are reviewed, including descriptions of fisheries, catch composition, catch rates and fisheries biology
studies conducted on target stocks. The most widely targeted coastal fish stocks are reef fishes and coastal pelagic fishes. Small pelagic species are
important for subsistence and smallscale commercial fisheries. Previously, small pelagic resources were important as a source of live bait for pole
andline tuna fishing, but this method is declining and only one large bait fishery is left in the region in the Solomon Islands. The poleandline bait
fisheries represent the only largescale industrial fisheries to have operated in the coral reef lagoons of the Pacific. Estuarine resources are of major
importance only in the large islands of Melanesia but are the staple diet of a relatively large proportion of the total South Pacific population. Deep
slope fish stocks form the basis of only two commercial fisheries in the region and expansion of deep slope fishing comparable with the 1970s and
1980s is unlikely to occur again. Commercial fisheries development is currently orientated towards small and mediumscale longline fisheries for
offshore pelagic resources, where high value tunas and billfishes are caught for export markets.
The total coastal fisheries production from the region amounts to just over 100000 tyr−1, worth a nominal US$ 262000000. About 80% of this
production is from subsistence fishing. Just under half the total annual commercial catch comes from fishing on coral reefs, which includes a small
tonnage of deep slope species. Invertebrates are the most valuable inshore fisheries resources and these include seacucumbers, trochus and pearl
oyster. Lobsters and mangrove crabs form the basis of smallscale commercial fisheries, as also do penaeid shrimps, except in PNG where they are
caught in large quantities through trawling. Mariculture of shrimps is becoming increasingly popular in the region and is a major industry in New
Caledonia. The greatest influence on coastal fisheries in the Pacific through the next decade is likely to come from southeast and east Asia, where the
demand for high value coastal fishes and invertebrates has led to large scale depletions and has motivated entrepreneurs to seek stocks in the
neighbouring Pacific islands.
Introduction
The archipelagos that are commonly referred to as the “South Pacific islands” are found in an area roughly bounded by the tropics and lying between 130°E and 125°
W (Fig. 1). There are three racial and cultural subgroupings of the South Pacific, namely Melanesia, Micronesia and Polynesia. The Melanesian islands are all relatively
large archipelagos and include Papua New Guinea (PNG), the Solomon Islands, Vanuatu and New Caledonia. Fiji lies halfway between Melanesia in the west and the
Polynesian islands of the central Pacific. Polynesia includes American Samoa, the Cook Islands, French Polynesia, Niue, Pitcairn, Tokelau, Tonga, Tuvalu, Wallis and
Futuna and Western Samoa. The Micronesian islands lie mainly north of the equator and include the Federated States of Micronesia—Yap (Fig. 2, 117)3, Chuuk (Fig.
2, 16), Pohnpei (Fig. 2, 79), Kosrae (Fig. 2, 43)—Guam, the Northern Mariana Islands, Marshall Islands, Nauru, Palau, and Kiribati.
3. Numbers in parentheses refer to specific locations within countries or territories and are indicated on Figure 2.
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Figure 1 Map of the South Pacific showing the countries and territories of the region. Easter Island (not shown) lies 2350km to the east
of Pitcairn Island.
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On the periphery of this grouping are the subtropical islands belonging to: Australia—Lord Howe (Fig. 2, 123), Norfolk Island (Fig. 2, 69); New Zealand—
Kermadec Islands (Fig. 2, 124); Chile—Easter Island, SalayGomez; the USA—Hawaii, Johnston Island (Fig. 2, 37), Wake Island; and Japan—Bonin (Fig. 2, 121),
Kazan Islands (Fig. 2, 122). The remaining islands comprise 15 independent countries, and seven territories belonging to either France, the USA or the UK.
Some basic geographic and economic information on the South Pacific island countries and territories is given in Table 1 (p. 409). Politically, these are all members
of the South Pacific Conference that was first convened in 1947 and which also includes the governments of New Zealand, Australia, the United States, Britain and
France. Fourteen independent island states, with the addition of New Zealand and Australia, are also members of the South Pacific Forum that was established in
1971. Both the Conference and the Forum have secretariats housed in New Caledonia and Fiji, respectively. Both institutions support fisheries development and
management in the South Pacific: the Forum through the Solomon Islandsbased Forum Fisheries Agency, which is concerned with managing access by distant water
fishing nations to the region’s tuna stocks, and the South Pacific Commission’s Fisheries Programme, which performs research and development on tuna and coastal
fisheries.
The combined area of the Exclusive Economic Zones (EEZs) of the South Pacific Commission island members (from the Northern Mariana Islands to the Pitcairn
Islands) is 29523000km2, but the land area of 550652km2 is a small portion of this. Most of the land (84%) belongs to PNG, with the other Melanesian islands and
Fiji forming a further 14% of the total. For many of the people of the South Pacific islands, particularly in landdeficient Micronesia and Polynesia, fish is a staple source
of animal protein. Although some islanders venture further offshore, fisheries in the coastal zone have traditionally been the target of subsistence activity and provide a
major portion of the diet. Even with increasing urbanization and the shift in preference to more imported western foods, fresh fish and invertebrates caught in coastal
waters continue to be a significant item in the diet of most Pacific islanders.
European exploration in the South Pacific during the last century and the Japanese entry during the 1920s and 1930s was followed by interest in the commercial
potential of invertebrate resources in the region, such as in molluscs for motherofpearl and pearls, and seacucumbers for bêchedemer production. Limited interest
was shown in the finfish stocks of the region, apart from some commercial exploitation of reef and tuna stocks in the Caroline Islands (Fig. 2, 14) during the 1930s.
Following the first World War, however, fishing for tunas such as skipjack (Katsuwonus pelamis), yellowfin (Thunnus albacares) and bigeye (Thunnus obesus),
continued to expand until at present the annual landings from the waters in the South Pacific Commission statistical area amount to about 9910001 worth in the region
of US$ 1460000000 (Anon. 1995). Following the Law of the Sea Conferences during the 1970s, the South Pacific islands claimed sovereignty over EEZs extending
200n.mi from the land. The present commercial catches of tunas are made predominantly within the EEZs of the Pacific islands by vessels from nations on the Pacific
rim such as Japan, Taiwan, USA, China, Korea and the Philippines (Anon. 1993a)
Impressive though the catches of tuna from the South Pacific are, they have limited impact on the lives of the indigenous peoples of the region. Tuna are caught by
highly mechanized industrial fleets of purse seiner, longline and poleandline vessels, often on the high seas. Less than 7% of this tuna is caught by Pacific island
vessels and only 25% of the total landings is processed within the region, at canneries in the Solomon Islands, Fiji and American Samoa. The remaining 75% of tuna
landings is processed elsewhere by countries on the Pacific rim (Anon. 1991a).
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Figure 2 Locations of places and geographical features in the South Pacific mentioned in the text. The first reference in the text to a
place or feature is followed by the corresponding number in parentheses.
Page 399
By contrast, landings from the coastal zone are more modest, but they have a far greater social and economic impact on the residents of the Pacific islands.
Moreover, there is a more immediate risk of overexploiting the resources in the narrow coastal zones of many Pacific island countries as populations increase and
technology improves the fishing power of artisanal fishers. Management of coastal fisheries is an increasingly important priority, but for this to have any hope of success,
information and feedback about the status and trends in coastal fisheries must first be acquired by fisheries managers and administrators. The objectives of this paper
are:
1. to describe the various coastal fisheries of the South Pacific,
2. review the various biological studies and stock assessment methods used to provide management information
3. to estimate the volume and value of these fisheries,
4. to discuss the possible future trends of these fisheries with respect to social, economic and political developments in the region.
In general we have restricted our summaries to the South Pacific islands, but refer where necessary to fisheries for the same or similar species on the periphery of the
region, particularly northeast Australia and Hawaii. Northeast Australia contains the Torres Straits (Fig. 2, 104) and shares several important fisheries resources with
Papua New Guinea. Hawaii, besides having strong social and cultural links with Polynesia, also provides a biological analogue for the subtropical islands in the south of
the region about which relatively little is known.
The first regional fisheries meeting convened by the South Pacific Commission (Anon. 1952) highlighted the lack of quantified information on South Pacific island
fisheries. However, because of the complexity and diffuse artisanal nature of coastal fisheries, together with the gradual development of national fishery administrations,
developments in fisheries production from the coastal zone have not yet been comprehensively documented at the regional level. The recent reviews of Pacific islands
fishery resources published by the Forum Fisheries Agency (Wright & Hill 1993) again highlighted the lack of information on the scale of harvests of fish and marine
organisms from the coastal zone. There is increasing concern about environmental issues such as sea level rise from global warming and loss of biodiversity through
excessive exploitation of living natural resources. Fisheries production may be affected by sea level rise and fisheries can contribute to local species depletions and
extinctions through excessive harvesting. This has already happened with the giant clam Tridacna gigas, which has been driven to extinction in many of the islands of
Micronesia, Vanuatu and probably New Caledonia (Munro 1993) and Fiji (Lewis et al. 1988a), and with certain reef fish species in parts of Micronesia (Myers 1989)
and Polynesia (Bell 1980, Hooper 1985, Sims 1990). These descriptions of fisheries and estimates of fisheries production are likely to be of interest to workers in a
variety of disciplines such as conservation, nutrition, economics, planning and coastal zone management.
The physical geography of the South Pacific islands

An inventory of the islands of the South Pacific, which includes notes on geology and structure of the different land masses in the region, is given by Douglas (1969).
Two basic types of island can be distinguished in the South Pacific, namely high islands and atolls. Land masses raised from the ocean floor through vulcanicity and
tectonic forces in time form high
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islands and develop fringing and coral reefs. Very young high volcanic islands such as Pagan in the Northern Mariana Islands have relatively little reef development
beyond encrusting coral communities. Older high islands may have well developed fringing reefs. Such is the case with Rarotonga (Fig. 2, 88) in the Cook Islands,
Tahiti (Fig. 2, 96) and Bora Bora in the Society Islands (Fig. 2, 94).
Nearly all the South Pacific islands lie within the tropics and so sea surface temperatures rarely fall much below 20°C and may rise as high as 30°C during the course
of a year. The coasts of most Pacific islands are characterized by coral reefs, seagrass meadows and mangrove forests. High islands contain the greatest number of reef
zones and habitats. They are also the only islands that have extensive fresh and brackish water habitats. Nutrientrich rivers may carry large quantities of silt resulting in
highly productive, but turbid muddy habitats. Mangrove forests thrive along the intertidal shorelines of estuaries and river mouths and seagrasses flourish on silty inner
reef flats and shallow lagoon floors.
High islands may subside, but the barrier and fringing reefs continue to grow and develop into atolls, where a fringe of coral islands and reef surrounds a lagoon.
Atolls lack rocky cliffs and platforms as well as rivers and the well developed mangrove communities found in high islands are either missing or poorly developed. They
therefore lack many of the species associated with these habitats. Occasionally, volcanic forces have raised atolls well above the sea surface to produce highly porous
limestone islands known as makateas. They lack rivers and have flat tops and steep sides that may plunge directly into the sea, be undercut or be fringed by rocky
platforms or reef flats. Some of the countries and territories of the South Pacific consist predominantly of only one island type such as atolls (Kiribati, Marshall Islands,
Tuvalu), makateas (Nauru, Niue) or high islands (Samoa, Vanuatu, Wallis and Futuna), but the remainder are usually a mixture of atolls, high islands and makateas.
Few parts of the South Pacific have such extensive freshwater discharge that coral reef development is inhibited over a wide area. The Gulf of Papua (Fig. 2, 32),
the region’s major estuarine area, has coral reefs at the western and eastern margins, where the influence of the massive freshwater influx from drainage of the
mountainous hinterland of Papua New Guinea is reduced. Elsewhere on the smaller islands of the Pacific, the outflow of rivers has a minor influence on reef
development.
The corals and coral reefs of the Pacific islands are described in Wells & Jenkins (1988) but the species of hermatypic or reef building corals in the South Pacific
have been fully described only for Australia (Veron 1986), with over 330 species contained in 70 genera. The number of species of coral declines in an easterly
direction across the Pacific in common with the distribution of fish and invertebrate species (see p. 404) so that there are only 30 genera present in the Society Islands
of French Polynesia and 10 genera in the Marquesa Islands (Fig. 2, 56) and the Pitcairn Islands. All forms of coral reef development can be found in the South Pacific
including large barrier reefs around New Caledonia and in Fiji, extensive fringing reefs, particularly around the large Melanesian islands, and patch and submerged reefs,
banks and shoals throughout the region.
Mangrove forests (especially Rhizophora spp., Bruguiera spp., and Avicennia spp.) are prevalent in estuarine areas but sediment build up may also permit
establishment of mangrove trees and bushes on the reef flat. On atolls, mangroves may be absent or present only in thin patches. A directory of Pacific island wetlands,
including mangrove forests, has been compiled by Scott (1993), while the distribution, environmental aspects and ecology of Pacific islands mangroves is reviewed by
Woodroffe (1987), and included in a global review of tropical marine ecosystems by Hatcher et al. (1989). Apart from the usefulness of the wood for building,
charcoal and tannin, mangrove forests act to stabilize areas where physi
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cal sedimentation is occurring and, from a fisheries perspective, are important as nursery grounds for peneaeid shrimps and some inshore fish species, and form the
habitat for some commercially valuable crustaceans. Extensive mangrove forests are a feature of high islands in the western Pacific, particularly the Melanesian islands
and Fiji. The natural eastern limit of mangroves in the Pacific is American Samoa, although Rhizophora stylosa was introduced to the Society Islands of French
Polynesia in the 1970s. Mangroves are also absent from Wallis and Futuna, Tokelau and the Phoenix (Fig. 2, 127) and Line Islands (Fig. 2, 128) of Kiribati.
Seagrasses are common in all marine ecosystems and are a regular feature of most of the inshore areas in the Pacific islands. According to Hatcher et al. (1989),
seagrasses stabilize sediments because leaves slow current flow, thus increasing sedimentation of particles. The roots and rhizomes form a complex matrix that binds
sediments and stops erosion. Seagrass beds are the habitat of certain commercially valuable shrimps, and provide food for reefassociated species such as
surgeonfishes (Acanthuridae) and rabbitfishes (Siganidae). Seagrasses are also important sources of nutrition for higher vertebrates such as dugongs and green turtles.
A concise summary of the seagrass species found in the western tropical South Pacific is given by Coles & Kuo (in press), and Wells & Jenkins (1988) include
information on seagrass beds in association with the coral reefs of the Pacific islands.
Climate and the marine environment of the South Pacific islands

Average annual rainfall in the South Pacific ranges from just over 1000mm in New Caledonia to 5000mm in Pohnpei and Kosrae States in the Federated States of
Micronesia. The large high island archipelagos of Melanesia may have quite different rainfall regimes on different parts of the same island or between locations on
different islands. For example, the rainfall in the Fijian capital Suva is about 3200mm per year while that of Nadi, some 110km to the west, is about 1900mm per year.
In PNG, Abeyasekera (1987) was able to distinguish three distinct rainfall regimes; namely, areas where rainfall is constant throughout the year, areas where rainfall
peaks between May and August and areas where rainfall is highest from December to March. High islands tend to retain moisture bearing clouds and have higher
annual rainfall regimes than atolls and other low islands. However, mountains may form rainshadow areas that receive rain only at certain times of the year as is the
case with the region around the PNG capital, Port Moresby (Fig. 2, 80), which experiences strong rainfall only during November to March and has an annual total of
about 1200mm per year.
For most of the Pacific islands rainfall typically ranges from 2000 to 3500mmyr−1. Low islands such as makateas and atolls tend to have less rainfall and may suffer
prolonged droughts. Furthermore, when rain does fall on coral islands and makateas where there is no major catchment area, there is little allochthonous nutrient input
into surrounding coastal waters and lagoons. Lagoons and embayments around high islands in the South Pacific are therefore likely to be more productive than atoll
lagoons. The productivity of highisland coastal waters, particularly where there are lagoons and sheltered waters, is possibly reflected in the greater abundance of small
pelagic fishes such as anchovies, sprats, sardines, scads, mackerels and fusiliers (Anon. 1984a). In addition, the range of different environments that can be found in the
immediate vicinity of the coasts of high islands also contributes to the greater range of biodiversity found in such locations.
Climatic seasonality in the South Pacific is more pronounced at higher latitudes. Even at
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or close to the equator there may be seasonal effects from the amount of rain carried by the prevailing winds. Most of the region is influenced by winds that blow from
the south and east (the Southeast Trades), but for about 4–5 months during the northern winter, the prevailing winds in the western Pacific blow from the north and
west (Northwest Monsoon). Rainfall tends to be highest during the summer and autumn months both north and south of the equator. This is well illustrated by
comparing the average monthly rainfall in Saipan (Fig. 2, 91) in the Northern Mariana Islands with that of Western Samoa. Both locations are at roughly the same
latitude north and south of the equator with the same average mean temperature (26°C), although Samoa is wetter with an average annual rainfall of 2900mm
compared with 2200mm in Saipan. Rainfall in Samoa reaches a maximum between November and February, while in Saipan rainfall peaks between July and October.
Information on the hydrographic characteristics of South Pacific marine environments has been summarized from various sources by Wauthy (1986). The waters that
form the surface layer of the tropical west and central Pacific enter into the transpacific intertropical circulation from the eastern boundaries of two subtropical
anticyclonic gyres, where the coastal upwelling of California and Peru provide enrichment of nutrient rich subsurface waters. The waters remain on the surface and the
thickness of the warm layer increases, isolating the surface waters from the deeper layers by a well established thermocline. As these waters move from east to west
they grow warmer and more impoverished as nutrients are consumed by photosynthesis and particulate materials are sedimented. Limited primary production continues
on the basis of partial remineralization within the isolated upper surface layer of the water column.
Nutrientdepletion leads to very clear blue oceanic water in which suspended particles are depleted and living organisms are scarce. The term “oceanic desert” has
been used by Lisitzin (in Wauthy 1986) to describe these nutrient poorwaters. Primary productivity in the photic zone ranges on average from 20 to 50gCm−2yr−1
(FAO 1972). Upwelling is one mechanism by which impoverished tropical waters can be enriched with nutrients from the subsurface waters and this has been
observed at the equator. Another mechanism whereby subsurface nutrientrich waters reach the euphotic zone involves shallowing of the thermocline at 10°N and 10°
S, at the edge of the equatorial counter currents. In the South Pacific, nutrient inputs from precipitation and runoff are of major significance only in the waters
surrounding the large island archipelagos of Melanesia where highlands are extensive and rainfall is very heavy. Not surprisingly, the highest oceanic primary
productivities in the region (90–180gCm−2yr−1) are found on the shelf area of the Gulf of Papua which receives much of the drainage from PNG highlands region.
Combination of various physical factors results in the accumulation in the tropical Pacific of a thick surface layer of warm water west of 180°. This accumulation
forms one of the preconditions necessary for the generation of cyclones or hurricanes that are a common meteorological phenomenon in the South Pacific. The second
precondition is the existence of a cycloniclike convergence in the lower layers of the atmosphere that can be found in the western tropical Pacific between the
equatorial monsoon winds from the west and the easterly trade winds. In the northwest tropical Pacific, cyclones form most frequently between June and November,
and are most frequent in August/September, with an average of 18 per year. South of the equator, cyclones occur from December to April and are less frequent than in
the northwest, with an average of four per year (Wauthy 1986).
Largescale oceanic events such as the El Niño Southern Oscillation (ENSO) also influence the coastal marine environment of the South Pacific islands. The
Southern Oscillation Index is the difference in atmospheric pressure between Tahiti and Darwin, which is usually
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positive due to the low pressure area over Indonesia and Australia. During an ENSO episode, the pressure gradient reverses and becomes negative for a prolonged
period with a consequent shift in climatic and oceanographic conditions. The easterly trade winds weaken and westerly winds are observed over parts of the equatorial
western Pacific. The area of warm water usually associated with the western tropical Pacific is displaced eastward over the central and eastern Pacific region and the
ocean waters of the western Pacific cool. This phenomenon results in the appearance of an anomalous warm ocean current off the coasts of Peru and Ecuador around
the Christmas season and hence was named by Peruvian fishermen “El Niño”, the familiar diminutive Spanish term for the infant Christ.
This major climatic shift produces unseasonal droughts in the western Pacific and unseasonal rains in the central and eastern Pacific. Information from commercial
tuna fisheries in the South Pacific and pelagic and demersal fisheries in South America suggests that ENSO events can, depending on species, have both negative and
positive effects on catchability and apparent abundance. In the western and tropical Pacific, the abundance of surface skipjack and yellowfin tuna stocks shifts
eastwards during an ENSO episode. This can be inferred from the concentration of fishing effort by tuna purseseine vessels, which during normal years concentrate to
the west of 160°E line of longitude and to the east of this line during an ENSO event (Anon. 1995). Little is known at present about how ENSO events affect coastal
fish and invertebrate stocks in the South Pacific due to the lack of any suitable time series of data. It is likely, however, that such a large scale anomaly will have an
influence on productivity and recruitment, especially in those species with long oceanic pelagic larval stages, and those reef species that are sensitive to anomalous water
levels during spawning or recruitment.
There may be other longterm climatic cycles in the Pacific region that will influence the productivity and abundance of marine organisms. Polovina et al. (1994)
describe such an event in the Hawaiian Islands that began in the mid 1970s and ended in the late 1980s. Over a 10year period, this climatic event promoted the
movement of nutrientrich deep ocean water into the euphotic zone during the first quarter of the year. This in turn resulted in higher survival of fish, crustaceans, seals
and sea birds. The decline in the event was followed by declines in the recruitment and abundance of fish, crustaceans, birds and seals. During this event an important
commercial lobster fishery in the Northwest Hawaiian Islands (Fig. 2, 72) expanded rapidly in the mid 1980s then declined as recruitment to the population was
markedly reduced, despite the efforts of fisheries managers to promote sustainable yields from the fishery.
Fisheries resources of the South Pacific

Marine resources in the context of this paper refer mainly to marine organisms that are caught and collected for food, but also include molluscs such as trochus, green
snail and pearl oysters where harvests are mainly for the shells. As the majority of the South Pacific islands are atolls and small islands surrounded by coral reefs, the
principal targets of nearshore fisheries in the region are fauna associated with coral reefs and lagoons. As stated previously, the only country with extensive estuaries is
PNG, whereas the other large Melanesian islands have smaller more limited estuarine environments. Species diversity of fishes, molluscs, crustaceans and echinoderms
declines in an easterly direction across the Pacific.
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There are about 2500 reef and inshore fishes in the Philippines, at the centre of the IndoPacific faunal continuum, compared with only 125 in Easter Island at the
eastern margin of the region (Myers 1989). This species gradient appears to be related to the position of the South Pacific islands in relation to the Pacific Plate, the
largest of the Earth’s lithospheric plates. The Pacific islands lie on or along the margin of this geological structure. The biogeography of the South Pacific region and
species distributions in relation to the Pacific Plate have been discussed by Springer (1982) and Myers (1989). Pacific islanders may consume a wide variety of reef
fishes, including snappers (Lutjanidae), emperors (Lethrinidae), groupers (Serranidae), parrotfishes (Scaridae), mullet (Mugilidae), surgeonfishes (Acanthuridae), jacks
(Carangidae), and other nearshore pelagic species such as scads (Carangidae), tunas and mackerels (Scombridae). Pacific islanders will also consume small species
such as squirrelfishes (Holocentridae), hawkfishes (Cirrhitidae) and some of the smaller surgeonfish species. Observations on a typical small scale commercial reef
fishery in the western and central part of the South Pacific may record between 200 and 300 species in the catch, although it is likely that only a few species will
dominate landings. The fishes commonly associated with mangrove and estuarine ecosystems in Melanesia are listed by Kailola & Wilson (1978), Collette (1983),
Quinn & Kojis (1986), Blaber & Milton (1990) and Thollot (1992). Species commonly caught in the large estuarine and mangrove areas include barramundi
(Centropomidae), catfishes (Ariidae), threadfins (Polynemidae), ponyfishes (Leiognathidae), clupeoids (Engraulidae & Clupeidae), jewfishes (Sciaenidae) and grunters
(Theraponidae).
South Pacific islanders also use a great variety of molluscs for food and for their shells. Cernhorsky (1967, 1972) records over 1000 species of shell bearing
molluscs from the South Pacific. In addition to these are the various cephalopods such as squids, cuttlefish and octopus that are caught in the nearshore zone. Several
molluscs are of prime commercial value in the region and these include trochus (Trochus niloticus), green snail (Turbo marmoratus) and blacklip pearl oyster
(Pinctada margaritifera). All these species are harvested primarily for motherofpearl used for button manufacture and furniture inlay. The blacklip pearl oyster, as
the name suggests, is also valuable for the production of a form of pearl that is dark silverygrey in colour and was originally collected from wild populations, but is
increasingly being cultured artificially. A wide variety of molluscs is also consumed for food and these are discussed in detail below.
There are an estimated 300 species of shallow water holothurians in the IndoPacific region that account for about 27% of the echinoderm fauna in the Pacific islands
(Guille et al. 1986). Holothurians form part of the subsistence diet of many Pacific islanders, although certain species are commercially valuable as a dried product
known as bêchedemer, or trepang, that is exported, mainly to Asia. There are at least 22 species of holothurians which are caught for bêchedemer production in
the South Pacific and these belong to the genera Actinopyga, Holothuria, Stichopus, Theloneta and Bohadschia (Preston 1993, Adams et al. in press).
Pacific islanders also consume a variety of crustaceans found in the nearshore zone including crabs, lobsters and shrimps. The mud crab (Scylla serrata) is widely
distributed in the region and this is caught for commercial sale as well as for subsistence. Other reef dwelling crabs such as the three spot reef crab (Carpilius
maculatus), the sand crab (Portunus pelagicus) and the red crab (Etisus splendidus) are also consumed for subsistence. Land crabs such as the coconut crab
(Birgus latro) have, traditionally been a component of subsistence catches and may be caught commercially, particularly where there is a developing tourist industry.
Other smaller land crabs such as Cardisoma carnifex and hermit crabs are a seasonally important subsistence resource.
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Several spiny lobster species are found in the South Pacific including Panulirus penicillatus, P. longipes, P. versicolor and P. ornatus, found mainly on tropical
reefs; and P. marginatus and P. pascuensis found on subtropical reefs. These and the related slipper lobsters (Scyllaridae) are captured both for subsistence and
commercial purposes. Other crustaceans that are harvested from the coastal zone include mantis shrimps (Squilla spp.), mud lobsters (Thalassina anomala) and
penaeid shrimps. Over 40 species of penaeid shrimps have been identified from the waters of PNG (Rapson & McIntosh 1972) but the most abundant is the banana
shrimp, Penaeus merguiensis. Also commonly captured are tiger shrimps, P. monodon and P. semisulcatus, and the endeavour shrimps, Metapenaeus ensis and M.
demani. Elsewhere, such as Fiji, Penaeus canaliculatus and Metapenaeus anchistus are locally abundant (Choy 1988), while species such as Penaeus
semisulcatus and Metapenaeus ensis, which are species of minor importance in PNG, are abundant in the lagoon of Tongatapu (Fig. 2, 103), the main island of
Tonga (Braley 1979).
Other invertebrates and marine organisms that are consumed regularly or as delicacies by Pacific islanders include chitons, seahares, marine worms and seaweeds.
Populations of the marine polychaete worm, Eunice viridis, (palolo in Samoan and balolo in Fijian) undergo periods of mass spawning in coastal waters once a year
during full moon periods. The gamete bearing segments of the worms rise to the surface where they can be collected by coastal villagers and are regarded as a great
delicacy in parts of the the Pacific, especially Samoa.
Fishing methods
Subsistence and artisanal fishing

There is a rich tradition of fishing techniques, beliefs and customs associated with fishing in the South Pacific islands and many of these have been described in
anthropological studies made over the last 200 years (e.g. Anell 1955). In this review, however, we are concerned mainly with contemporary fishing practices and will
describe those gears and fishing methods that are widely used on a regular basis in the region. Most coastal fisheries in the South Pacific are characterized by small
scale artisanal fishing methods. A considerable amount of fishing takes place from the shore or in shallow waters without the use of fishing vessels. Where fishing vessels
are used, these are generally small, either nonpowered canoes or canoes and dinghies powered by outboard motors and, to a lesser extent, by sail. Larger vessels of
8–20m in length, powered by inboard diesel engines, are used for commercial fishing for demersal species beyond the reef slope, and for catching tuna on the open
ocean.
Common gears include hooksandlines, traps (fixed and moveable), seine and gill nets, and spears. Hooksandlines can be deployed in a variety of ways, as
simple droplines to catch demersal fishes, as bottom and surface longlines to catch demersal and pelagic fishes respectively, and towed with baits and lures to troll for
pelagic fishes. Traditionally, hooks were fashioned from shells, bones and wood, whereas lines were made from coconut or other plant fibre. These traditional materials
have generally been superseded by monofilament lines and metal J or circle hooks. Traditional shell lures are still used in some locations such as French Polynesia
(Chapman & Cusack 1988a) to troll for tuna and other large pelagic species. Handline or dropline fishing in shallow coastal waters is a common subsistence and
recreational fishing method in most of the Pacific. Handlines can be deployed
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on reefs, in estuaries and on the shelf to catch demersal species. Handlines are also used in midwater to catch small pelagic fishes such as bigeye scads or round scads,
using baits or lures. Coastal fishermen will also use handlines in midwater to catch tuna and other large pelagic species such as rainbow runner (Elagatis bipinnulatus)
and wahoo (Acanthocybium solandri).
Commercial droplines for demersal species such as snappers and groupers on the deep reef slope or on banks and seamounts are mounted on reels to aid hauling
from depths between 100 and 400m. A common design of handreel for such operations was developed in Western Samoa by the Food and Agricultural Organisation
of the United Nations and propagated throughout the region by the South Pacific Commission (Dalzell & Preston 1992). Longline fishing has also been used to catch
demersal species from the deep reef slopes, particularly in Fiji, where longlines of between 500 and 1000 hooks were set on offshore banks and seamounts. Similar
sized surface longlines are presently employed to catch tuna, particularly large yellowfin and big eye tunas that have a high value on overseas markets in Japan and
Hawaii.
Gill netting, beach seining and drivein netting are conducted both in coralline and estuarine areas of the Pacific. Nets were traditionally manufactured by Pacific
islanders from plant fibres such as coconut and pandanus, but such nets are now rarely made and used except in the most remote islands. Gill net fishing is practised on
reefs and lagoons in the Pacific and in some areas, such as Kiribati, has become one of the most popular fishing methods in this archipelago. Gill net fishing is also
widely practised in estuarine areas of the Pacific. In most instances nets are set from dinghies and canoes for periods of between 1h and an overnight soak. The major
drawback for fishermen with gill nets in both reefs and estuaries is damage to the nets from sharks.
Drivein net fishing is commonly practised around many islands of the Pacific. Nets are set in an area of shallow water, on a reef plateau or in the lagoon and fish are
driven by scare lines and swimmers into the net. The fish may be concentrated at one end of the net for hauling, or swimmers armed with spears may enter to kill and
collect fish. A description of this type of fishing operation is given in Smith & Dalzell (1993). Surround netting involves setting a net around an area of coral or around a
school of fish. The fish are then caught by swimmers who enter the net enclosure carrying spears. Beach seines may be deployed in lagoon waters to trap schooling
fishes such as scads, small jacks, herrings and halfbeaks, while barrier nets can be strung across reef passages and channels to trap fish as they return from feeding on
the reef plateau.
Other common net fishing techniques include cast netting and scoop netting for flying fishes. Cast nets are used in coastal shallows by fishermen to catch schooling
species such as mullet and rabbitfishes. The fishermen stalk the school, which often creates a ripple pattern on the water surface, and attempt to cast a circular weighted
net over the school and thus entangle the fish in the mesh. Handheld scoop nets are commonly used in Polynesia to catch flying fishes at night. Fishermen chase the
flying fishes over the water surface, spotting them by torch light and catching them in the scoop nets before the fish can launch themselves into flight (Gillett & Ianelli
1991).
Spears may be single or multiplepronged, traditionally made from wood and bone but nowadays made of steel. Spear fishing is conducted both above water and
below. Spear fishermen may target fish from land or boat using spears and arrows, or by diving beneath the water with hand spears and spear guns. Captured fish are
threaded on a line wrapped around the diver’s waist, on a line trailing behind the diver or even towed in a galvanized basin buoyed by an old car innertube, as a
precaution against shark attack. The development of
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masks, fins, SCUBA gear, steel spears and spear guns has meant that the fishing power of the spearfishermen has greatly increased. Some spear fisheries are very
specialized such as that for dolphinfish (Coryphaena hippurus) in French Polynesia. Highpowered launches will chase the dolphinfish along the surface, as this species
will usually not dive to escape pursuit. As the fish tires the fisherman harpoons it with a multiplepronged barbed spear.
Fixed or stationary traps are a common feature in coastal areas of the South Pacific. The simplest of these structures are Vshaped stone and stick enclosures with
an entrance that faces the shore, as found, for example, in PNG (Hulo 1984) and Cook Islands (Baquie 1977). They may be more complex structures comprising a
series of leaders or barriers that guide the fish into a series of interconnecting chambers. The chambers terminate in a single catching chamber where the fishes may be
netted or speared. These more complex structures are found in French Polynesia (Grand 1985), Guam (Amesbury et al. 1986), Tonga (Halapua 1982) and Palau
(Johannes 1981). Fixed barrier traps take advantage of the tidal foraging migration of different species of fish to effect capture. Fishes that have been feeding on the
reef flat or in estuarine shallows will follow the receding tide into deeper water. When they encounter a fence they will swim along it and concentrate in a chamber or net
where they can be caught.
The regular use of portable fish traps appears to be confined mainly to Micronesia and parts of French Polynesia, although bamboo and mangrove wood traps were
traditionally deployed in coastal areas of the South Pacific (see for example Koch 1961 and contributions in Quinn et al. 1984). Johannes (1981) describes the
deployment of portable fish traps in the shallow coastal waters of Palau, and Smith & Dalzell (1993) give a brief account of trap fishing in Woleai Atoll (Fig. 2, 115),
which lies to the east of Yap. Traps in Palau are made either of traditional materials, such as sticks and vines, or welded steel bars and chicken wire and are used to
catch a variety of reef fish. Traditional stick and vine traps are also used on Woleai to catch reef fishes and one type of trap is specifically designed to catch the goatfish
(Mulloides flavolineatus) when seasonally abundant in the lagoon. Cubiform wire mesh traps are deployed in the lagoon of Rangiroa Atoll (Fig. 2, 88), Tuamotu
Archipelago in French Polynesia, to catch surgeonfishes, especially Acanthurus xanthopterus and A.bleekeri.
A variety of other fishing and collecting activities are conducted along Pacific shorelines and reefs in addition to fishing with spears, lines, traps and nets. Kite fishing
is still employed in some parts of the region to catch needle fish (Belonidae) (Johannes 1981, Hulo 1984). A spider web lure is towed behind a canoe beneath a
Pandanusleaf kite. Needle fish or longtoms, which prey mainly on small pelagic fish, will attack the web lure as it skips over the sea surface and become snared as
their teeth tangle in the spider web. Molluscs and echinoderms can be picked off the reef at low tide, and octopus may be drawn out of holes in the reef with a metal
hook. In most locations molluscs, crustaceans, seacucumbers and seaurchins, collected mainly by women and children, may form a significant fraction of the total reef
harvest (Wass 1982, Mathews & Oiterong 1991, Rawlinson et al. 1994).
Larger predatory fishes may also be caught in nooses at the water surface. Migrant phosphate mine workers from Tuvalu and Kiribati have been observed to noose
wahoo (Acanthocybium solandri) that aggregate around mooring buoys on Nauru (Cusack 1987). A bamboo pole is rigged with a short line and a teaser bait, usually
a flying fish, attached to one end. A second pole has a running noose attached to one end. The bait is splashed on the water surface until a wahoo responds and begins
to make passes at it. The fisherman then attempts to position the noose between the wahoo and the bait so that the fish will pass through and can be snared. Perhaps
the most spectacular example of this type of fishing is the catching of reef sharks by fishermen from Kontu village on New Ireland (Fig. 2, 67),
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PNG (Kohnke 1974). Reef sharks are attracted or called by the use of a coconut shell rattle shaken in the water, which draws the shark to the canoe. The shark is
coaxed through a cane noose with a reef fish (preferably rabbitfish). The noose is attached to a wooden propeller that spins as the shark dives, tightening the cane
around the gills and suffocating it.
Industrial scale fisheries

Most of the islands of the South Pacific have steeply shelving slopes and are surrounded by coral, unsuitable for conducting trawl fishing. Trawls can be deployed on
the soft bottoms of estuarine areas of the bigger islands, however, for catching shrimps and demersal fishes. Only one South Pacific country, PNG, has established
commercial trawl fisheries. The most important of these is the trawler fleet in the Gulf of Papua that numbers between 10 and 20 vessels and targets stocks of penaeid
shrimps. The trawlers displace about 150 gross tonnes (gr.t.) and are twin rigged with two 12fathom (footrope) nets. This fleet fishes 24h per day throughout the year
in the northern and eastern Gulf of Papua and has been operational since 1969. In the early 1980s, a smaller shrimp fishery, limited in most years to two vessels,
commenced in Orangerie Bay (Fig. 2, 74) on the southeast Papuan coast.
The lagoons and sheltered embayments of the Pacific islands often contain small pelagic fishes that can be used as live bait for poleandline tuna fishing. The small
pelagic resource is usually a complex of clupeoid species including anchovies (Engraulidae) and sprats, sardines and herrings (Clupeidae) as well as small scads
(Carangidae), mackerel (Scombridae), silversides (Atherinidae) and fusiliers (Caesionidae). The small gracile anchovies belonging to the genera Encrasicholina and
Stolephoms, and the sprats of the genus Spratelloides, are the principal targets of tuna livebait fisheries in the South Pacific.
Fleets of poleandline tuna vessels (mainly from Okinawa) have operated from Palau, PNG, Kiribati, Fiji, New Caledonia and the Solomon Islands. A typical
vessel is about 20m in length and displaces about 60gr.t. Catches of live bait are made from the poleandline boats using a stickheld dip net or boukeami. Catches
are made at night after concentrating the bait fishes around powerful submersible lights. The captured bait is concentrated in one end of the net and loaded into buckets
for transfer to tanks set in the deck of the tuna vessel. The boukeami is then stowed while the tuna vessel sails for the open sea to pursue tuna schools. The
deployment and operation of the boukeami is described in detail by Muyard (1980). Poleandline tuna fishing has largely been superseded by purseseining and is
now confined chiefly to the Solomon Islands and Fiji fleets with small fisheries in Tuvalu and Kiribati (Anon. 1995).
Finfish fisheries
Reef fisheries
Description
Fishing on coral reefs occurs in all the countries and territories of the South Pacific and most of the techniques employed are comparable between the different
locations. We have therefore not described at length the reef fisheries in each country; instead we have summarized the relevant literature in Table 1 and selected
examples that might be described as typical of
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a country or subregion within the South Pacific. Without exception, all reef fisheries in the South Pacific are smallscale fisheries using simple nonmechanized gears.
The major differences between Pacific island reef fisheries are the range of gears used, the emphasis on particular gears and most importantly, the fishing pressure
applied to reef fish stocks. Fish does not appear to be as important in subsistence diets in the large island archipelagos of Melanesia as in the smaller resourcelimited
countries of Micronesia and Polynesia (Coyne et al. 1984). The annual per capita production for the Pacific islands can be approximated from the figures in Table 27
(p. 499) and the population figures in Table 1. This ranges from 7–40kg or a mean of 23kg for the Melanesian islands, while for the Polynesian and Micronesian islands
the ranges are 6–121kg and 4–170kg, with means of 61kg and 63kg respectively. Fishing effort on all coastal fisheries stocks, including reef fishes, is substantially
greater in the Micronesian and Polynesian islands, in comparison with the Melanesian islands.
In all but the remotest islands, reef and other coastal fisheries have both a subsistence and a commercial component. The size of the commercial fisheries sector is
dependent on the degree of urbanization of the island population, the amount of available land for agriculture and the development of the cash economy. Subsistence
reef fishing methods tend to be very simple and in most of the examples given in Table 2, fishes caught by handlines and spears make up most of the subsistence catch.
Larger volumes of fish can be generated through the use of community fishing methods such as group spearing and drivein net fishing and these methods may be
employed when greater amounts of fish are required for occasions such as village feasts. Commercial reef fisheries also include handlines and spears but in the
examples in Table 2 there is a greater emphasis on the use of nets such as gill nets, drivein nets and fish corrals.
Table 1 Geographical and economic statistics for the countries and territories of the South Pacific.
Country Land area (km2) Population (1994) Pop. density Annual growth rate (%) Total GDP ($US 000) Per capita GDP
American Samoa 200 54600 273 3.7 203125.3 5194.8
Cook Islands 237 19100 81 1.1 70095.5 4052.1
Federated States of Micronesia (FSM) 701 105900 151 3.0 246011.2 2652
Fiji 18272 777700 43 2.0 1,620707.4 2118.5
French Polynesia 3521 218000 62 2.5 3,202764.2 15305.2
Guam 541 146700 271 2.3 1,180427.8 9637.7
Kiribati 810 78300 97 2.3 33875.4 468
Marshall Islands 181 54069 299 4.0 74735.8 1556
Nauru 21 10600 505 2.9 160875 17486
Niue 259 2100 8 −2.4 6891.6 3077.8
Northern Mariana Is. 471 56600 120 9.5 571297 10094
New Caledonia 19103 182200 10 2.0 2,125919.6 12753
Palau 488 16500 34 2.2 49367.1 3247.4
Papua New Guinea 462243 3,951500 9 2.3 5,670260.7 1468
Pitcairn 5 60 12
Solomon Islands 27556 367400 13 3.4 262526.2 738.7
Tokelau 10 1500 150 −1.3 624 372.8
Tonga 747 98300 132 0.5 138035 1415.7
Tuvalu 26 9500 365 1.7 64187.2 7053.5
Vanuatu 12190 164100 13 2.8 208878.5 1308.8
Wallis & Futuna 255 14400 56 1.3
Western Samoa 2935 163500 56 0.5 165885.7 1017.9
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Table 2 Summary of literature sources describing reef fisheries in the Pacific islands.
Country Location Reference

Australia Torres Straits Johannes & MacFarlane (eds) 1991
American Samoa Tutuila Craig et al. 1993, Saucerman 1994
Cook Islands Rarotonga Baquie 1977
Palmerston Anon. 1988a
Federated States of Micronesia Smith 1992a
Woleai Atoll Smith & Dalzell 1993
Kosrae Wilson & Hamilton 1992
Yap Uwate 1987
Fiji Rotuma Anon. 1983a
Rabi (Fig. 2, 84) Anon. 1983b
Dravuni (Fig. 2, 20) Emery & Winterbottom 1991,
South & East Fiji Jennings & Polunin 1995a,
Viti Levu Rawlinson et al. 1994
French Polynesia Tuamotu Archipelago Grand 1983, 1985, Stein 1988
Kaukura Grand 1983
Rangiroa Grand et al. 1983
Tikehau Morize 1985, Caillart 1988a
Guam Amesbury et al. 1986, Katnick 1982,
Knudson 1987, Myers 1993
Hawaii Smith 1994
Kiribati Gilbert Islands Mees et al. 1988
Tarawa Yeeting & Wright 1989
Marshall Islands Smith 1992b
Nauru Dalzell & Debao 1994
New Caledonia Loubens 1978a, Leblic & Teulieres
1987, Anon. 1988b, 1994a
Niue Dalzell et al. 1992
Northern Mariana Islands Hamm et al. 1994
Palau Johannes 1981 1991, Nichols 1991,
Mathews & Oiterong 1991, Kitalong & Dalzell 1994
Papua New Guinea Kavieng Wright & Richards 1985, Dalzell & Wright 1990
Port Moresby Lock 1986a,b,c,d
Western Manus Chapau & Lokani 1986
Pitcairn Islands Pitcairn Island Sharpies 1994
Solomon Islands Marovo Lagoon Hviding 1988
Roviana, Tulagi and Vona Vona Leqata et al. 1990, Blaber et al. 1990
Gizo Sasabule 1991
Roviana GinaWhewell 1994
Ontong Java BaylissSmith 1974
Tokelau Fakaofo (Fig. 2, 23) Hooper 1985
Tonga Tongatapu Zann 1981, Halapua 1982, Munro 1990
Tuvalu Funafuti Zann 1980, Patiale & Dalzell 1990
Nanumea Chambers 1984
Vanuatu David & Cillauren 1989
Wallis & Futuna Wallis Taumaia & Cusack 1988
Futuna (Fig. 2, 27) Galzin 1985
Western Samoa Upolu Helm 1992, Zann et al. 1991
In PNG, the largest of the Melanesian archipelagos, fishing activity, including subsistence fishing, is very limited around most of the coast and this is true of fishing on
coral reefs (Dalzell & Wright 1986). Wright & Richard’s (1985) study of the Tigak Islands (Fig. 2, 100) reef fishery in northern PNG is a good descriptive account of
subsistence and commercial reef fishing applicable to most coral reef areas of PNG and probably most of rural Melanesia. Almost all the subsistence catch came from
handline fishing and spear fishing from
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canoes and dinghies. Wright & Richards note that catches are made to satisfy only immediate food requirements and nets are rarely employed as their use requires
more organization and yields a large surplus catch. There is no fulltime commercial reef fishery in the Tigak Islands. When cash is required, villagers will occasionally
catch relatively large volumes of fish for sale in the main urban centre of Kavieng (Fig. 2, 39), mostly through gill and surround netting. The main influences on the
volume of commercial landings by individual villages were distance from markets in Kavieng and the price of the main agricultural commodity, copra (Dalzell & Wright
1990).
The main motivation for commercial fishing in the Melanesian islands is lack of any agricultural land. Daugo Island (Fig. 2, 19) is a small barren coral island on the
Papuan Barrier Reef with poor top soil, few trees and bushes and no fresh water, which the islanders have to bring from the mainland. As the Daugo Islanders have no
subsistence or cash crops they have turned to fulltime professional fishing and are among the main suppliers of fresh reef fish to Port Moresby, the capital city of PNG.
Lock (1986a,b,c,d) described the Daugo Island reef fishery where twothirds of the catches come from surround nets, drivein nets and gill nets deployed from
outboardpowered dinghies. The balance of the catch comes from handlining, spear fishing and trolling. Unlike the villagers of the Tigak Islands who fish only
occasionally, the fishermen of Daugo Island are active daily, fishing from dawn until midafternoon, when they travel to the produce markets in Port Moresby to sell
their catches.
Descriptions of reef fishing for subsistence purposes in the Solomon Islands (BaylissSmith 1974, Hviding 1988, Leqata et al. 1990) and Vanuatu (David &
Cillauren 1989) are comparable with those in PNG. Subsistence catches are taken mainly with handlines from canoes. Sales of copra and vegetable produce are
among the main sources of income in most locations, rather than commercial fishing. As with PNG, the most successful commercial reef fishermen are landless migrants
from other provinces who have no alternative incomes from agriculture (Sasabule 1991). Both New Caledonia and Fiji have large urban populations and well
established commercial reef fisheries, both of which are dominated by landings of emperors, particularly Lethrinus nebulosus (Dalzell et al. 1992, Anon. 1994a).
Commercial reef fish catches in both countries are taken mainly with handlines and gill nets (Loubens 1978a, Anon. 1988b, Rawlinson et al. 1994). New Caledonia
and Fiji, in particular, have well developed tourist industries with over 80000 and 280000 visitors per annum, respectively, that provide an extra market for commercial
reef fish catches.
Subsistence reef and lagoon catches in the rural areas in both New Caledonia and Fiji and are taken by a combination of spear fishing, handlining and net fishing
(Leblic & Teulieres 1987, Emery & Winterbottom 1991, Rawlinson et al. 1994, Jennings & Polunin 1995a, b). In Fiji, around 30% of rural Fijians go fishing at least
once a week (37% of the males, 48% of the females, and 5% of the children, on the main island of Viti Levu), but this involves nearly 100% of village households
(Rawlinson et al. 1994). The main reef fish targeted are species in the family Lethrinidae, but serranids, hemiramphids, gerreids and scombrids are also prominent, and
carangids are important around the larger islands. Fiji has approximately 2000 registered artisanal fishing boats (Anon. 1994b). New Caledonia has a large fleet of
recreational craft, over 11000 registered vessels mostly berthed in the capital city, Noumea (Fig 2, 71) (NguyenKhoa 1993). According to Loubens (1978a), about
60% of the total catch from the southwest lagoon of New Caledonia is taken by recreational fishermen, using mainly handlines and spear fishing. Furthermore Kulbicki
& Grandperrin (1988) showed that there was a positive correlation between catch per unit effort (CPUE) of lagoon species and distance from Noumea in a 50km
radius after which catch rates level off. The
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50km radius is approximately the outer limit of the range of most recreational fishermen (Loubens 1978a).
As fish, in general, comprise a greater component of the diets of Polynesians and Micronesians, this demand creates more intensive subsistence and commercial
fisheries than in Melanesia. In Tonga and Western Samoa, nearly 70% of the respective populations are concentrated on one island in each archipelago, namely
Tongatapu and Upolu (Fig 2, 109). Zann et al. (1991) state that over three quarters of the villages on Upolu are engaged in fishing with an average of between two
and four fishing trips per week and that half the households in rural areas are reliant on reef fisheries as the main subsistence source of protein. Almost all fishing occurs
in shallow inshore waters less than 10m deep, with spear fishing being the most regularly employed fishing method, followed by net fishing and handlining. Two thirds
of the catch is consumed with the balance being sold in markets in Apia (Fig 2, 4), or increasingly sent by air to American Samoa, where there is a very high demand
for fresh reef fish (Craig et al. 1993).
Tongan reef fisheries have been described by Halapua (1982) and more recently by Tu’avao et al. (1994). Commercial reef fish catches are made by spear fishing,
handline fishing, gill and drivein netting and from fish corrals. The commercial reef fisheries in Tonga are structured around smallboat operators who employ a
number of fishermen and take the major share of the catch revenue. Spear fishing and net fishing are conducted on the reefs around Tongatapu; however, handline
fishermen have been forced to travel to offshore reefs and banks due to depletion of stocks around the main island. Fish corrals set on the reef plateau around
Tongatapu were mainly constructed for catching mullet, and were so successful that stocks declined markedly, leading to a ban (not fully implemented, however) on this
type of gear. The decline in catches has led to a decline in the number of fish corrals around Tongatapu. Landlessness is also a factor in the commercial reef fishery in
Tongatapu. Most of the spear fishermen in the commercial fishery are migrants from the Ha’apai Islands (Fig 2, 33) who, possessing no land, turn to a simple and
cheap fishing method to provide incomes.
Fishes are one of the very few resources that offer any commercial potential in the Polynesian and Micronesian atolls, and reef and lagoon fishes are the easiest and
most convenient for fishermen to target. Commercial reef fisheries in French Polynesia supply the main island of Tahiti where half of the population of this territory now
live. About 40% of the fish sold through the market in Tahiti are reef fishes from the atolls of the Tuamotu Archipelago (Fig 2, 107). Descriptions of the reef fisheries in
these atolls have been given by Grand (1983, 1985), Grand et al. (1983), Morize (1985) and Caillart (1988a). About 90% of the commercial reef fish landings in these
islands are from fish corrals deployed on the atolls of Kaukura (Fig 2, 38), Aratua (Fig 2, 5), Tikehau (Fig 2, 100), Rangiroa and Apataki (Fig 2, 3) (Stein 1988).
Other fishing techniques such as gill nets, handlines, portable fish traps and spear fishing are used to supplement commercial catches, when production from the corrals
is reduced. Similar movement of commercial reef fish catches from an outlying atoll to the main urban centre is also found in the neighbouring Cook Islands. Relatively
large volumes of parrotfishes, mullet and trevallies, caught with gill nets and drivein nets at Palmerston Atoll (Fig 2, 78), are regularly sent to markets in Rarotonga
(Richards 1993).
Reef fishing in the remoter atolls of Micronesia and Polynesia are entirely for subsistence, apart from the small amounts of fish that might occasionally be traded with
passing ships. Smith & Dalzell (1993) described reef fishing methods used on Woleai Atoll, one of the outlying atolls in the Caroline Islands, where many aspects of
traditional life are maintained. Reef fishing methods include handlines, spear fishing, portable fish traps and drive
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in net fishing. While this is similar to other islands in the Pacific, gears like leaf sweeps used for driveinfishing and fish traps are still manufactured from traditional
materials. Furthermore, techniques such as fish trapping require considerable skill and there are several different trap types deployed in different seasons to target
various reef fish species. Increasing urbanization in atolls, as elsewhere, is leading to changes in reef fisheries with the development of commercial fisheries and the
adoption of more modern efficient fishing techniques. At Tarawa (Fig. 2, 98), the principal atoll of Kiribati and home to 40% of the population of the Gilbert Islands
(Fig. 2, 30), the numbers of monofilament handlines and gill nets increased exponentially over a 10year period to become the most important fishing methods for reef
and lagoon fishes (Yeeting & Wright 1989).
Many of the American associated islands of Micronesia (Palau, Guam, Northern Mariana Islands, Federated States of Micronesia) are becoming increasingly
urbanized as populations grow and accelerate demand for fresh fish and in particular fresh reef fish. Tourism, mainly from Japan, is a major industry in islands such as
Guam, Palau and Saipan, and is also creating an additional demand for reef fish. A large range of traditional techniques to catch reef fish have been documented on
Guam (Amesbury et al. 1986) but, as elsewhere, they are largely replaced with more modern gears. Commercial reef fishing on Guam comprises mainly handline
fishing and spear fishing, with minor contributions from various net fishing methods (Myers 1993). Commercial handline fishing targets reef fish stocks in waters
<150m, mainly emperors (Lethrinidae), jacks (Carangidae) and snappers (Lutjanidae). Spear fishing consists of two components based on whether SCUBA
equipment is used. Higher proportions of certain species such as wrasses (Labridae), groupers (Serranidae), parrotfishes (Scaridae) and goatfishes (Mullidae) are
caught by spear fishermen using SCUBA equipment. Commercial spear fishermen on Guam have driven certain reef species, such as the bumpheaded parrotfish
(Bolbometopon muricatum) and the Napoleon wrasse (Cheilinus undulatus), to very low population levels and similar practices are thought to be depleting
populations of the same species on Palau (Myers 1989, Hensley & Sherwood 1993).
Traditional and contemporary fishing methods for catching reef fish in Palau, include handlining, spear fishing, gill netting, fish trapping and set nets (Johannes 1981).
Women and children on Palau obtain a subsistence catch of fish primarily by handlining in reef pools during the day or spearing fishes from the surface at night
(Mathews & Oiterong 1991). No information is available on the relative contribution to commercial and subsistence catches by the various gears, although Johannes
(1981) states that set nets and spear fishing are the two most important and productive reef fishing methods employed on Palau. Set nets, known locally as kesokes,
work on the same principle as fish corrals in that they trap fish as they migrate into deeper water with the falling tide, and concentrate the catch in the apex of a
stationary Vshaped barrier net (Johannes 1981). Besides markets in the capital, Koror (Fig. 2, 41), reef fishes are also exported to Guam and Saipan, both as official
exports and in the form of passenger luggage on planes. An increasing number of fish buyers are coming to Palau from Guam and the Northern Mariana Islands to buy
fish for sale at home and transporting it in ice boxes as luggage (Preston 1990a, Adams 1993a), which means that this is not registered in the official exports from
Palau.
Reef fisheries in countries with very limited reef and lagoon areas such as Niue, the Northern Mariana Islands and Pitcairn Island may still form an important
component of total fisheries landings. In the Northern Mariana Islands, surrounded mainly by narrow fringing reefs and with limited lagoon area, reef fishes account for
half of commercial fish landings. The principle fishing methods include gill nets, fish fences or corrals, handlining and spear fishing (Watt 1990). Nearly all subsistence
catches are shallow water reef fishes taken by
Page 414
similar gears. Niue is a makatea or raised atoll with a narrow coralencrusted rock shelf. Half of the total fish catch on Niue comes from fishing on the narrow reef
around the island, and about twothirds of this catch is taken by handline fishing (Dalzell et al. 1993). Commercial fishermen on Niue concentrate on deep slope and
pelagic fish and all reef fish catches are part of the subsistence production. The small population on Pitcairn Island fish regularly for subsistence and, perhaps surprisingly
for such a remote location, for commercial purposes (P. Sharpies, South Pacific Commission, pers. comm.). Because of historical interest in Pitcairn, however, several
vessels stop by the island each month for the Pitcairners to market fish and carvings. Fishing is conducted on the fringing reefs and reef slope around Pitcairn Island,
either from dinghies or from the rocky shore. Almost all fishing is conducted with handlines targeting the drummer, Kyphosus bigibbus, and the grouper, Epinephelus
fasciatus, for subsistence, while the red grouper, Variola louti, and the larger eteline snappers on the deep slope are the targets of commercial fishing.
By contrast, fisheries production on Nauru and Futuna, both islands with narrow fringing reefs and no lagoon area, is mainly from pelagic rather than reef fisheries.
Nauru, like Niue, is a makatea but it is unusual in that despite its small size it is a very wealthy country, as a result of substantial revenues generated from mining the
phosphate bearing rock that forms most of the interior of the island. Nauruans fish mainly for recreational purposes from powered skiffs, trolling for large pelagic
species rather than fishing for reef fishes (Dalzell & Debao 1994). The several thousand Kiribati and Tuvaluan mine workers on Nauru supplement their incomes by
commercial fishing, mainly for large pelagic fishes. A little commercial handline fishing for reef fishes is conducted, however, on the reef slope in waters between 50 to
100m, often using a Tshaped or cruciform wire assembly, known as a Christmas Tree, with between 18 and 32 hooks. Small amounts of reef fish are also caught
commercially by spear fishing using SCUBA. In all, reef fishes comprise about 10% of the total fish landings on Nauru. Reef fishes are also caught on Futuna for
subsistence, using handlines, spears, and gill nets (Galzin 1985). As on Nauru reef, however fishes make only a minor contribution (≈6.0%) to the total fisheries
production on the island, most of which comes from exploitation of large pelagic fishes.
The foregoing summarizes the salient features of reef fisheries in the South Pacific islands. We will also briefly mention the characteristics of reef fisheries on the
Australian Great Barrier Reef (GBR) (Fig. 2, 29) and around the Hawaiian archipelago. Commercial reef fishing on the GBR is highly specialized and mainly targets the
coral trout, Plectropomus leopardus, and the emperor, Lethrinus chrysostomus, together with other high value snappers, groupers and emperors. Catches in both
the commercial and recreational fisheries are made with handlines. Other reef fishes that are highly valued by Pacific islanders such as surgeonfishes, parrotfishes,
rabbitfishes and wrasses are generally not targeted by commercial and recreational fishermen. The total annual commercial reef fish catch from the GBR amounts to
about 3000t (Brown et al. 1994). Reef fisheries in Hawaii exhibit more similarities to those in the Pacific islands, although subsistence catches are not as important and
recreational catches form the largest proportion of the noncommercial catch. Like New Caledonia, Hawaii has a large fleet of recreational craft, estimated to be
12690 vessels and nearly 190000 anglers (Smith 1993). Reef fishes form a relatively small part of the total commercial inshore catch of about 700tyr−1 (≈10%) but
market preferences increase the value of the reef fishes catch (Smith 1993). Reef fish are caught predominantly with handlines and gill nets by commercial fishermen,
while angling with rodandreel assemblies, and spear fishing with and without SCUBA equipment generate most of the recreational reef fish catch.
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Catch composition
The composition of reef fish catches is extremely varied, both in time and location. Table 3 presents information on the composition of reef fish landings from various
locations in the South Pacific classified to the family level. The dominant feature of fish landings in many parts of the Pacific is the emperors (Lethrinidae), while other
important components include surgeonfishes (Acanthuridae), snappers (Lutjanidae), parrotfishes (Scaridae), and coastal tunas and mackerels (Scombridae). Although
these families contain a large range of species, often one or a few species will dominate in each family. In New Caledonia and Fiji, for example, landings of emperors
consist mainly of Lethrinus nebulosus (Dalzell et al. 1992, Anon. 1994b), while in Tikehau Atoll (French Polynesia) emperor landings consist mainly of Lethrinus
miniatus (Caillart 1988a). Similarly, surgeonfish landings in New Caledonia are dominated by Naso unicornis (Anon. 1992a), while Acanthurus dussumieri and
Naso spp. (mainly N. unicornis and N. brevirostris) constitute most of the surgeonfish landings in the reef fishery on the South Papuan coast (Lock 1986a). Although
commercial reef fish landings in Palau contain over 200 species, half of the catch comprises only seven species (N. unicornis, Bolbometopon muricatum, Lutjanus
gibbus, Lethrinus ramak, Hipposcarus longiceps, Siganus canaliculatus, S. lineatus).
Pelagic species such as the small mackerels, Rastrelliger spp., and the large predatory Spanish mackerel (Scomberomorus commerson) are common components
of reef fish landings although not reef fish sensu stricto. Fishermen trolling along the reef edge often catch large pelagic predators such as Spanish mackerels, dogtooth
tunas (Gymnosarda unicolor), rainbow runners and queenfish (Scomberoides spp.), which may be included in the reef fish catch. Furthermore, landings of carangids
may be dominated by the small bigeye scads, Selar crumenophthalmus, and the less common S. boops. These small schooling pelagic fishes are universally prized as
a food fish throughout the Pacific and in some years, when particularly abundant, will dominate landings of jacks and of reef fishes in general (Helm 1992, Saucerman
1994). A feature of bigeye scad landings in reef fisheries is their inconsistency, where they can dominate landings in one year and be virtually absent from the fishery the
next (Helm 1992, Hensley & Sherwood 1993, Saucerman 1994)
Catch composition will also depend on the type of fishing gear employed in the fishery. Handlines catch predominantly predatory species such as snappers,
groupers and emperors, but small hooks baited with items such as seaweed, coconut flesh and congealed squid ink can be used to catch herbivores such as
surgeonfishes, rabbitfishes and rudderfishes (Amesbury et al. 1986). Spear and net fishing tend to take a broader range of species. This is well illustrated by Table 4,
which shows the catch composition for three different methods of net fishing plus handlining and spear fishing on shallow reef stocks of the South Papuan Barrier Reef
near Port Moresby. Twothirds of the handline catch are emperors with most of the balance of the catch made up from snappers, jacks and groupers. Emperors are
also a dominant feature of the net catches, but there is a wider range of species taken including herbivores such as surgeonfishes, parrotfishes and rabbitfishes. Spear
fishermen rely on getting close to their quarry and emperors tend to shy away from divers underwater. Other species, such as surgeonfishes, groupers and scombrids
such as Spanish mackerel, that become curious of divers, are vulnerable to spear fishermen.
There are few long time series of catch species composition data for reef fisheries in the South Pacific from which to judge the longterm effects of fishing on reef fish
communities. Kitalong & Dalzell (1994) examined a 14yr time series of commercial reef fish landings data from Palau and concluded that there had been little change
in the compostion of landings over time at the level of the family. Declines were noted, however, of certain species, such
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Table 3 Percent composition of reef fish catches from various locations in the South Pacific region.
Family Fiji PNG PNG Solomon Kiribati Palau Guam N. Mariana Kosrae Am. W. Tonga Tikehau (Fr. Nauru New
(North) (South) Is Islands1 (FSM) Samoa Samoa Polynesia) Caledonia
Lethrinidae 17.66 10 29 6.2 7.8 14 17.70 1.35 5.36 1.44 19.5 38.36 6 31.02
Chanidae 0.32 2
Balistidae 0.33 2 0.57 0.97 2 4.74
Albulidae 0.01 1 16.5 2
Hemiramphidae 1.66 1
Serranidae 12.62 9 3 6.6 3.1 9 7.58 1.01 5.21 12.84 4.55 4 3.62 18
Scombridae 18.74 3 10 15.3 24.3 1.39 12.57
Carangidae 7.41 14 8 10.2 3.2 4 16.59 1.26 7.88 8.05 1.59 18 3.26
Mullidae 1.48 5 5.1 2.9 1 1.81 0.38 3.56 2.22 8.05 4.56 10 0.77
Lutjanidae 11.01 13 5 19.2 11.1 12 3.63 0.26 7.36 7.02 1.75 1.33 6 39.54 3.50
Acanthuridae 4.06 5 7 6.4 14 9.12 1.11 22.73 27.76 19.5 16 20.78 3.65
Scaridae 3.84 8 5 11.5 18 8.48 5.55 6.91 6.33 14 2.69 10 1.17
Belonidae 0.17 5 3.9 0.33 0.02 0.61
Mugilidae 10.24 21 4 3.3 1 0.36 0.04 6.57 7.47 10.45 17.00 6 1.12 10.85
Siganidae 1.08 1 6 3.9 10 5.62 1.80 7.31 0.01 1.05 1.43 2 3.91
Sphyraenidae 7.45 3.2 7.90 0.10 1.07 2 0.24
Gerridae 0.21 2 5.3 0.01 0.07
Haemulidae 1.33 3 4 5.4 0.01
Labridae 4.31 0.03 2.77 0.8 0.16
Holocentridae 5.05 12.47 2.25 4.47 17.26
Theraponidae 0.39 0.03
Others 8 6 6.3 19.3 17 15.99 87.10 22.06 18.62 10.05 26.12 16 6.62 16.51
1. Most reef fish landed in the Northern Mariana Islands are reported only as mixed reef fish.
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Table 4 Percent catch composition by different artisanal fishing methods on reef fish stocks on the South Papuan Barrier Reef.
Fishing method
Family Spear Hand line Gill net Drivein net Surround net
Carcharhinidae 1 2
Mylobatidae 1
Chanidae 2 1
Hemiramphidae 3
Belonidae 0 1
Serranidae 21 10 3 1
Carangidae 5 7 11 13 4
Lutjanidae 3 12 13 6
Gerridae 0 3
Haemulidae 14 10 5 1
Lethrinidae 1 65 42 31 37
Sparidae 0
Mullidae 3 12
Mugilidae 11
Platacidae 4
Kyphosidae 8 2 1
Sphyraenidae 2 1
Labridae 2 2
Scaridae 3 1 7 10
Acanthuridae 17 2 2 17
Siganidae 0 8 5
Scombridae 17 11 5
Balistidae 3 0 1
Others 3 3 2 6 0
Total 100 100 100 100 100
as Crenimugil crenilabis, that were thought to be linked to excessive fishing pressure. Stein (1988) summarized information from a 25yr time series of landings of reef
fishes from the Tuamotu Archipelago but, unfortunately, did not comment on whether there have been major shifts in the catch composition over this time. Catch data
between 1982 and 1991 from Guam were analyzed by Hensley & Sherwood (1993) who showed that there were large declines of certain species in the reef fish catch
such as goatfish (Mulloides flavolineatus), parrotfish (Bolbometopon muricatum), wrasse (Cheilinus undulatus), and large snappers and groupers.
Catch rates
Catch rates from deployment of artisanal gears such as handlines, spears and nets are at best modest (Tables 4–7). Catches from handline fishing in shallow water
ranged from 0.03 to 12.12kgline−1h−1 and from 0.44 to 3.5kgline−1h−1 for overall mean catch rates. In most instances, the handline gear consists of monofilament line
equipped with between 1 and 3 hooks, usually with some form of sinker, although in Nauru, wire rigs bearing between 18 and 32 hooks (see p. 414) are preferred to
maximize catches. The principal targets of shallow water handline fishing are groupers, snappers, emperors and jacks. The highest catch rates in Table 5, reported by
Dalzell & Preston (1992), are from fishing mainly on snapper and grouper stocks beyond the reef margin in waters between 80 and 400m in depth (see p. 432). Catch
rates from underwater spear fishing ranged from 0.08 to 9.6kgman−1h−1, with
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Table 5 Summary of catch rate (CPUE, catch per unit effort) and catch composition data for handline fishing on shallow coral reef and deepslope fish
stocks in the South Pacific.
CPUE
(kgline−1h−1)
Location Target stock range mean Principal components Source
Papua New Guinea (Port Shallow reef species 0.68–4.0 2.46 Lutjanidae, Carangidae, Serranidae, Lock 1986a,c
Moresby) Lethrinidae
Papua New Guinea (Tigak Is) Shallow reef species na 1.2 Lutjanidae, Carangidae, Serranidae, Wright & Richards 1985
Lethrinidae
American Samoa Shallow reef species 0.25–1.51 0.54 na Saucerman 1994
Guam Shallow reef species 0.03–2.04 0.55 Carangidae, Lethrinidae, Acanthuridae, Katnik 1982
Siganidae
Palau Shallow reef species 2.2–7.32 3.49 Lutjanidae, Lethrinidae, Serranidae Anon. 1992e 1993b
Fiji (Ba Fig. 2,6) Shallow reef species 0.14–12.12 2.27 Lutjanidae, Lethrinidae, Carangidae, J.Anderson, MRAG, London, pers.
Serranidae comm.
FSM (Kosrae) Shallow reef species na 1.78 na Smith 1992a
Tonga (Tongatapu) Shallow reef species na 0.44 Lethrinidae, Holocentridae, Lutjanidae Munro 1990
Tuvalu (Funafuti) Shallow reef species 0.33–5.93 2.35 Lutjanidae, Lethrinidae, Serranidae, Patiale & Dalzell 1990
Carangidae
Wallis Shallow reef species na 1.3 Lutjanidae, Lethrinidae, Serranidae, Taumaia & Cusack 1988
Carangidae
FSM (Pohnpei) Shallow reef and deep slope 0.69–5.12 3.01 Lutjanidae, Carangidae, Serranidae, Dalzell unpub. data
species Lethrinidae
FSM (Yap) Shallow reef and deep slope 0.97–3.1 1.67 na Uwate 1987
species
FSM (Chuuk Outer Banks) Shallow reef and deep slope 1.31–4.57 2.30 Lutjanidae, Carangidae, Lethrinidae, Diplock & Dalzell 1991
species Scombridae
Nauru Shallow reef and deep slope 0.75–7.2 3.0 Lutjanidae, Serranidae, Carangidae, Dalzell & Debao 1994
species Holocentridae
Tropical Pacific atolls Deep slope species 0.4–19.0 7.7 Lutjanidae, Serranidae, Carangidae, Dalzell & Preston 1992
Lethrinidae
Tropical Pacific high islands Deep slope species 2.2–13.2 6.0 Lutjanidae, Serranidae, Carangidae, Dalzell & Preston 1992
Lethrinidae
Tonga Deep slope species 3.64–5.31 4.5 Lutjanidae, Serranidae, Lethrinidae Mees 1994
Papua New Guinea (Kavieng) Deep slope species 0.6–11.24 3.1 Lutjanidae, Serranidae, Lethrinidae Richards & Sundberg 1984
Papua New Guinea Deep slope species na 3.7 Lutjanidae, Serranidae, Carangidae, Chapau 1988
Lethrinidae
FSM (Pohnpei) Deep slope species 3.9–5.5 4.7 Lutjanidae, Carangidae, Serranidae, McCoy 1990
Lethrinidae
Tuvalu Deep slope species 1.72–9.62 6.1 Lutjanidae, Serranidae, Carangidae Anon. 1993c
Vanuatu Deep slope species 0–5.00 1.44 Lutjanidae, Serranidae, Lethrinidae Schaan et al. 1987
Niue Deep slope species 2.1–8.5 5.5 Lutjanidae, Serranidae, Carangidae Dalzell et al. 1992
na=not available
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Table 6 Summary of catch rates (CPUE, catch per unit effort) and catch composition of spearfishing on South Pacific reefs.
CPUE (kg
man−1h−1)
Location Target stock range mean Principal catch components Source
Papua New Guinea (Port Reef & lagoon species & large 1.2–3.6 2.4 Serranidae, Acanthuridae, Scombridae, Lock 1986a
Moresby) pelagics Haemulidae
Papua New Guinea (Kavieng) Reef & lagoon species na 2.4 Scaridae, Serranidae, Lutjanidae, Wright & Richards 1985
Haemulidae
Palau Reef & lagoon species 7.4–9.6 8.5 Scaridae, Serranidae, Acanthuridae, Anon. 1992e, 1993b
Lethrinidae
Nauru Reef and reef slope species 0.2–4.3 2.0 Lutjanidae, Holocentridae, Serranidae, Dalzell & Debao 1994
Acanthuridae
Guam Reef & lagoon species 0.08–1.14 0.41 Scaridae, Kyphosidae, Siganidae, Katnik 1982
Acanthuridae
Kosrae Reef & lagoon species 1.8–5.7 3.28 na Smith 1992a
Woleai (Micronesia) Reef & lagoon species 0.55–2.04 1.27 Acanthuridae, Scaridae, Balistidae, Labridae Smith & Dalzell 1993
Tonga Reef & lagoon species 1.2–1.7 1.4 Acanthuridae, Scaridae, Holocentridae, Halapua 1982
Serranidae
Fiji (Dravuni) Reef & lagoon species 0.81–1.6 1.20 Serranidae, Acanthuridae, Lutjanidae, Emery & Winterbottom 1991
Carangidae
Fiji (Ba) Reef & lagoon species 0.12–5.7 1.51 Lethrinidae, Lutjanidae, Serranidae, J.Anderson, MRAG, London
Scombridae (pers.comm.)
na=not available
averages ranging from 0.4 to 8.5kgman−1h−1. The principal targets for spear fishing are groupers, surgeonfishes, parrotfishes and snappers (Table 6).
Tables 7 and 8 give details of catches by gill nets, drivein nets and beach seining in the Pacific islands. Unlike spear fishing and handlining where CPUE is invariably
reported as kg man−1h−1 or kgline−1h−1, a range of different units to express CPUE have been used. Comparisons are further complicated by the variation in net length
and mesh sizes employed. Catch per set is the commonest reported CPUE for gill nets and this ranged from 4.2 to 31.8 kg set−1 for the examples given here. The
principal targets for gill net fishing appear to be jacks, mullets, emperors, goatfishes and snappers. The same problems in the expression of effort and CPUE were
found with the examples of drivein net fishing in Table 8. Catch rates for driveinnet fishing varied between 0.25 and 3.9kgman−1h−1, or 13.1 and 42.7kgset−1. The
beach seine catches in Table 8 were all directed towards catching small pelagic fishes in coral reef lagoons, and all but one of these were taken from the published
records of the South Pacific Commission’s Skipjack Survey and Assessment Programme. The very high average catch rate for beach seining in Rabaul (Fig. 2, 83),
PNG was a result of regular fishing on a large school of bigeye scads (Selar spp.). Elsewhere, in the Pacific, beach seine catches, predominantly of small pelagic fishes,
tended to be lower, with catches comprising sardines, herrings, goatfishes and jacks.
Records of catch rates from traditionally manufactured portable fish traps set on reefs in the South Pacific could not be found in the literature. Available information
on performance of fish traps comes from experimental deployment of Caribbeanstyle traps on Pacific reefs to catch species in the shallow reef community or on the
deep reef slope to catch mainly large groupers and snappers. Dalzell (1996) quotes average catch rates of between 0.85 and 4.6kgtrap−1 for traps set on shallow reefs
and 3.2 to 8.9kgtrap−1 for traps set on deep reef slopes. Caillart & Morize (1985) quote average catch rates from fish corrals at Tikehau Atoll
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Table 7 Summary of catch rate (CPUE, catch per unit effort) and catch composition data for gill net fishing on reef and small pelagic fish stocks in the
South Pacific region
Location Net length Mesh size Target stock CPUE Principal catch components Source
(m) (cm) range mean
American Samoa Reef & lagoon 3.3–6.8kgh −1
5.0kgh −1 na Saucerman 1994
species
Kiribati na 5.7–12.7 Reef & lagoon 5.0–96.0kgtrip−1 43.4kgtrip−1 Albulidae, Carangidae, Mugilidae, Anon. 1991c
species Mullidae
Solomon Islands na 5–15 Reef & lagoon 0.26–0.90 kg 0.46kg 100m Sharks, Chanidae, Carangidae, Blaber et al. 1990
species 100mnet−1h−1 net−1h−1 Mugilidae
Cook Islands 90–230 4.5–5.0 Small pelagics & 0.14–18.04 kg 2.2kg 10mnet−1 Carangidae, Priacanthidae, Chapman & Cusack
reef fish 10mnet−1 Mullidae, Caesionidae 1988d
Fiji (Rabi Island) 150 1.9–7.6 Reef & lagoon 15–26kgset−1 18.9kgset−1 Lethrinidae, Lutjanidae, Mugilidae, Anon. 1983a
species Holocentridae
Fiji(Rotuma Fig. 2, 90) 229 7.6 Reef & lagoon 10.0–60.0kgset−1 31.8kgset−1 Mugilidae, Carangidae, Lutjanidae, Anon. 1983b
species Lethrinidae
Papua New Guinea na 5.0–12.7 Reef & lagoon na 2.0kgman−1h−1 Lethrinidae, Lutjanidae, Lock 1986a
(Port Moresby) species Carangidae, Scombridae
Papua New Guinea 35–100 3.8 Small pelagics 0.7–6.7kgset−1 3.0kgset−1 Carangidae, Clupeidae Dalzell 1993a
(Rabaul)
Papua New Guinea 100 7.6–15.0 Reef & small 0.0–17.2kgset−1 7.57kgset−1 Sharks, Scombridae, Chanidae, Chapau & Lockani
(Manus) pelagics Clupeidae 1986
Kosrae(FSM) na na Reef & lagoon 4.2–9.1kgh−1 6.3kgh−1 na Smith 1992a
species
Tonga 100–1200 5.0 Reef & lagoon 5.6–7.2 kgset−1 6.0kgset−1 Acanthuridae, Labridae, Halapua 1982
species Siganidae, Lethrinidae
Guam na na Reef & lagoon 0.67–12.24kgset−1 4.24kgset−1 Acanthuridae, Mullidae, Scaridae, Katnik 1982
species Labridae
na=not available
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Table 8 Summary of catch rate (CPUE, catch per unit effort) and catch composition data for driveinnet and beach seine fishing on reef and small
pelagic stocks in the South Pacific region
Location Net length Mesh size Target stock CPUE Principal catch components Source
(m) (cm) range mean
Drive in net
Woleia (Micronesia) 35 4.5 Reef & lagoon 8.1–129.4kgset−1 42.7kgset−1 Acanthuridae, Scaridae, Smith & Dalzell 1993
species Siganidae, Lethrinidae
Palau na na Reef and small 2.3– 5.1kgman−1h−1 Carangidae, Lethrinidae, Anon. 1993b
pelagic species 11.1kgman−1h−1 Acanthuridae, Siganidae
Papua New Guinea (Port na 5.0–12.7 Reef & lagoon 1.41– 2.52kgman−1h−1 Lethrinidae, Carangidae, Lock 1986a
Moresby) species 4.95kgman−1h−1 Mugilidae, Siganidae
Papua New Guinea (Tigak 100 6.3–7.5 Reef & lagoon na 3.9kgman−1h−1 Mugilidae, Chanidae, Wright & Richards
Is) species Carangidae, Scaridae 1985
Papua New Guinea (Manus) 100 7.6 Reef & lagoon 0.45– 0.69kgman−1h−1 Scaridae, Mugilidae, Chapau & Lokani
species 0.74kgman−1hr−1 Carangidae, Acanthuridae 1986
Cook Islands (Palmerston 14–480 2.3 Reef & lagoon 0–41kgset−1 13–1kgset−1 Scaridae Anon. 1988a
Atoll) species
Nauru na na Reef & lagoon 1.1–8.0kghr−1 3.9kghr−1 Kyphosidae, Mugilidae, Dalzell & Debao 1994
species Acanthuridae, Lutjanidae
Guam 140–280 na Reef & lagoon 0.09– 0.25kgman−1h−1 Siganidae, Acanthuridae, Katnik 1982,
species 0.46kgman−1h−1 Labridae, Lethrinidae Amesbury et al. 1986
Beach seine
Papua New Guinea (Rabaul) 200 2.5 Small pelagic na 350kgset−1 Carangidae, Clupeidae Dalzell 1993a
species
Papua New Guinea 100 2.5 Small pelacic 1.0–44.6kgset−1 17.3kgset−1 Carangidae Dalzell unpub. data
(Kavieng) species
Butaritari (Kiribati) (beach 80 0.5 Small pelagic 8–345kgset−1 130.6kgset−1 Clupeidae, Atherinidae, Kleiber & Kearney
seine) species Apogonidae 1983
French Polynesia 80 0.5 Small pelagic 5–33kgset−1 18.6kgset−1 Clupeidae, Mullidae, Carangidae Gillett & Kearney
(Marquesas Is) species 1983
Tokelau (Fakaofo Atoll) 80 0.5 Small pelagic 7.5–25.5kgset−1 13.5kgset−1 Mugilidae, Clupeidae SSAP 1983a
species
Tonga (Vava’u) 80 0.5 Small pelagic 13–18kgset−1 15.0kgset−1 Atherinidae, Clupeidae, SSAP 1983b
species Mullidae
Marshall Islands (Majuro & 80 0.5 Small pelagic 16–38kgset−1 21.5kgset−1 Clupeidae, Atherinidae, SSAP 1984
Jaluit Atolls (51)) species Lethrinidae
na=not available
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during the peak of the fishing season (NovemberFebruary) of 380–580kgday−1. Grand (1983) does not give daily catch rates for fish corrals at Kaukura Atoll but
reports that the average fish production ranged from 1 to 2.3tcorral−1month−1, with a mean of 1.5tcorral−1 month−1.
As stated earlier a variety of other methods are used to catch fishes on coral reefs, although documentation on these is scarce. Average catch rates for cast netting on
reefs in Nauru (Dalzell & Debao 1994), Guam (Katnick 1982) and Kosrae (Smith 1992a) were 0.7kgman−1h−1, 2.8kgman−1h−1 and 5.6kgman−1h−1 respectively, and
comprised rudderfishes (Kyphosidae), rabbitfishes, surgeonfishes, jacks and goatfishes. Collection of invertebrates and molluscs is a common pastime on reefs for
women and children, and may add significantly to the overall harvest from the reef (see p. 427). Mathews & Oiterong (1991) made a detailed study of women’s reef
fishing activities and found that they target over 25 species of echinoderms, molluscs and crustaceans for collection, as well as catching fish with handlines and nets.
Catch or harvest rates of all species for gleaners in Kosrae ranged from 2.2 to 4.1kgh−1, with a mean of 3.6kgh−1.
Fisheries biology and stock assessment

The biology of reef fishes has received considerable interest in the past 50 yr, due in part to the development of SCUBA equipment and the ability of biologists to
conduct observations in situ. Sale (1980) provided the first major review of reef fish biology and ecology, and more recently various contributions in Sale (1991) have
provided the most comprehensive review of various aspects of reef fish biology. Aspects of the biology of South Pacific reef fishes are also covered by Wright (1993)
and Pyle (1993) who, together with Sale, include information on the biology of reef fishes in the Atlantic and Indian Oceans. Many of the studies on reef fish biology in
the South Pacific have been conducted on the Australian GBR, Hawaii, French Polynesia and Guam. The focus of much of the research on Pacific reef fishes in the last
five decades has been on small strongly siteassociated species, such as damselfishes, with which it is relatively easy to conduct underwater observations and
experiments. However, Munro & Williams (1985) pointed out in their review of reef fisheries management that there is a general dearth of stock assessment studies of
commonly exploited reef fish populations, either in the South Pacific or elsewhere, and that there are very few estimates of life history parameters and population studies
such as age, growth, mortality, and recruitment that are essential for stock assessment.
We shall not undertake a detailed biological review in this section, nor in the subsequent sections on other coastal fisheries resources. We are mainly concerned here
with documenting those studies that have been conducted in the South Pacific on the most commonly exploited fish and invertebrate stocks, and which have generated
information on life histories relevant for management. From Table 3 (p. 416), the principal reef fish families include the emperors, surgeonfishes, snappers, jacks,
parrotfishes, groupers, mullet, goatfishes, rabbitfishes and squirrelfishes. As stated earlier, coastal scombrids may form an important component of reef fish catches, but
the biology of these species in the Pacific is reviewed further below (p. 444). We will also review the methods used to estimate reef fish standing stocks and to compute
maximum sustainable yields (MSYs) from reef fish stocks in the region and compare yields among the different locations in the Pacific.
The emperors or lethrinids are probably the most widely studied of the Pacific reef fishes that form major components of reef and lagoon fishery landings. Lethrinids
have proved very amenable to ageing using well established techniques for reading annuli in otoliths and
Page 423
scales, as well as through more recent techniques for ageing using otolith microstructure (daily increments) and length based methods. Loubens (1978b, 1980)
estimated age and growth of emperors and a range of other reef fish species from otolith annuli including the economically important species Lethrinus nebulosus and
L. miniatus (=L. chrysostomus). Age and growth of L. nebulosus from the Australian GBR have also been determined through reading of otolith annuli (McPherson
et al. 1985). Scales (Walker 1975) and otoliths (Brown et al. 1994) have both been used to age L. miniatus from the GBR and subsequent agefrequency
distributions were used to generate catch curves for mortality estimation. Otolith microstructure or daily growth increments were also used to estimate the age and
growth of L. olivaceus (=L. miniatus) in French Polynesia (Caillart et al. 1986), L. semicinctus in northern PNG (Mobiha 1993), and L. rubrioperculatus in
American Samoa (Ralston & Williams 1988a). Age, growth and mortality of exploited stocks of emperors, mainly L. nebulosus, L. mahsena and L. harak, have been
studied in Fiji (Dalzell et al. 1992) using a combination of length based methods and age estimates from otoliths. L. harak was also included in a study of commonly
exploited reef stocks in Palau (Kitalong & Dalzell 1994), where growth and mortality estimates were generated from lengthfrequency data.
Life spans of the larger Lethrinus species such as L. nebulosus and L. olivaceus would appear to be in excess of 20 yr, while small species such as L.
nematacanthus and L. semicinctus have life spans of between 7 and 10 yr. Loubens (1978b, 1980) also estimated age and growth from otolith annuli for the smaller
lethrinids or breams in New Caledonia belonging to the genus Gymnocranius, including G. japonicus, G. lethrinoides and G. rivulatus. Age and growth of another
bream, Nemipterus peroni (=furcosus) (Nemipteridae), was also included in Loubens’ (1980) study and Chapau (1993) has used otolith annuli and tagging to
estimate age, growth mortality rates and abundance of the same species in populations in northern PNG. Both groups of breams have average life spans of about 5–10
yr.
Surgeonfish biology and life histories have been studied in a number of locations in the South Pacific, most notably Acanthurus triostegus in Hawaii (Randall 1961),
Naso brevirostris in Tikehau Atoll (French Polynesia) (Caillart 1988b) and Acanthurus nigricauda and A. xanthopterus in northern PNG (Dalzell 1989). Age and
growth of these species have been described from a combination of observations on captive specimens (A. triostegus), otolith microstructure (A. nigricauda, Naso
brevirostris), length frequency data (Acanthurus xanthopterus, Naso brevirostris) and tagging (Acanthurus triostegus, A. xanthopterus). Daily growth increment
formation was validated in the otoliths of several juvenile surgeonfishes including Ctenochaetus binotatus, C. striatus, Zebrasoma scopas and Z. veliferum from the
GBR (Lou & Moltschanowskyj 1992). Otolith microstructure has also been used to provide preliminary age and growth estimates for Acanthurus lineatus and
Ctenochaetus striatus from American Samoa (Ralston & Williams 1988a). C. striatus has been studied in some detail in French Polynesia as it is frequently
implicated in cases of ciguatera poisoning (Bagnis 1970). Attempts to estimate growth for this species from tagging data were unsuccessful (Walters 1968, Bagnis
1970) but growth, mortality and recruitment were determined for C. striatus from Moorea using lengthfrequency data (AriasGonzalez et al. 1993). Similar analyses
were also conducted with lengthfrequency data for Naso unicornis from Palau (Kitalong & Dalzell 1994).
Surgeonfishes appear to have relatively long life spans. Randall (1961) reported adult surgeonfishes such as Naso unicornis and Acanthurus xanthopterus living
for between 15 and 20yr in captivity. Age and growth studies of A. nigrofuscus populations on the GBR suggest that this species, one of the smallest acanthurids, has
an average maximum life span of over 20yr (Hart & Russ in press), together with another common species, Ctenochaetus striatus
Page 424
(N.V.C.Polunin & E.D.Brothers unpub. data). Furthermore, 40 presumptive annuli have been observed in the otoliths of Acanthurus lineatus from the same location
(A.Fowler, Bedford Institute of Oceanography, pers. comm.; Polunin & Brothers unpubl. data). It should be noted that surgeonfishes on the GBR are subject to
virtually no fishing pressure and populations only experience mortality through natural causes.
Some aspects of the biology of the rabbitfishes or siganids have been studied both in wild populations and from studies in captive populations in Guam (Tsuda et al.
1976). The species covered in the various contributions in Tsuda et al. (1976) include Siganus spinus, S. rostratus and S. argentus. The biology of wild populations
of S. lineatus and S. canaliculatus was observed from wild populations in Palau (Drew 1973), while in Fiji the biology of S. vermiculatus was described from a
combination of observations on wild and captive populations (Gundermann et al. 1983). Only the growth of juveniles was observed in each of these various studies,
either in captive populations (Tsuda et al. 1976, Gundermann et al. 1983) or from tagging of juveniles (Drew 1973). The biology of S. canaliculatus in Palau has also
been described by Hasse et al. (1977), while the population biology of this and other rabbitfish species (S. argentus and S. lineatus) in Palau were described by
Kitalong & Dalzell (1994) from length frequency data.
The biology of shallow water snappers has received less attention than the more commercially valuable deep slope species. Loubens (1978b, 1980) succeeded in
ageing in New Caledonia Lutjanus amabilis, L. bohar, L. fulviflamma, L. kasmira, L. quinquelineatus and L. vitta by reading otolith annuli. In the same location,
Baillon & Kulbicki (1988) have aged the sweetlip, Diagramma pictum, from otolith and scale annuli and from otolith microstructure. Although not strictly a snapper,
this species belongs to the family Haemulidae, which is closely related to the Lutjanidae. Wright et al. (1986) described the biology of the red bass, Lutjanus bohar, in
northern PNG and estimated age, growth and mortality parameters from lengthfrequency data. Lengthfrequency data were also used by Kitalong & Dalzell (1994) to
generate the same parameters for L. gibbus in Palau.
Ralston & Williams (1988a) included the small bluelined snapper, L. kasmira, in studies of depth distributions, growth and mortality of deep slope fishes in the
Northern Mariana Islands. Age estimates in this study were made from otolith microstructure, whereas mortality rates were computed from length data. The age and
growth of L. kasmira in American Samoa have also been estimated from otolith microstructure, while MoralesNin & Ralston (1990) estimated the age and growth of
the same species in Hawaii, where it is an introduced exotic, from both otolith annuli and daily increments. Another small snapper, L. fulvus, from French Polynesia
(Caillart et al. 1986) was aged using otolith microstructure. Age and growth of three large snappers from the Australian GBR, L. sebae, L. malabaricus and L.
erythopterus, were determined from otolith annuli (McPherson & Squire 1992). Although these three species are not strictly speaking shallow water snappers, they
can be found in a greater range of depths than the eteline snappers, including shallow reef areas. Longevities of between 10 and 20yr appear to be typical for Lutjanus
spp. and Loubens (1980) reports life spans in excess of 20yr for even small species such as L. fulviflamma and L. quinquelineatus.
Parrotfishes, although a common component of reef catches, have received little attention from fisheries biologists in reef areas. The formation of annular marks in the
otoliths of Scarus schlegeli from the GBR has been validated by Lou (1992), while daily increments in the otoliths of a range of juvenile parrotfishes, including S.
rivulatus, S. globiceps, S. psittacus, S. sordidus, S. niger, S. frenatus and S. oviceps (all from the GBR) have been observed by Lou & Moltschaniwskyj (1992)
and Lou (1993). Coutures (1994) used annular marks on scales to age the largest of the parrotfishes, Bolbometopon muricatum, in New
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Caledonia, and estimated mortality rates from lengthfrequency data. Kitalong & Dalzell (1994) also used lengthfrequency data to generate growth and mortality
parameters for the same species in Palau. Lou’s (1992) estimates of age and growth of Scarus schlegeli, one of the species in the middle of the size range of the
Scaridae, suggest longevities of between 6 and 9yr, while Coutures’ data for Bolbometopon muricatum indicates a life span of about 25 yr. Polunin & Brothers
(unpubl. data) using a combination of scanning electron and light microscopy on putative annual and finer banding in otoliths, inferred longevities of over 20yr for Scarus
globiceps, S. sordidus and S. frenatus. The related wrasses or Labridae, appear to have received even less attention than the parrotfishes. Kitalong & Dalzell (1994)
estimated growth and mortality parameters from lengthfrequency data for the large Napoleon wrasse, Cheilinus undulatus, in Palau. Preliminary studies on this
species from the GBR suggest that ageing through reading otolith annuli is practicable and that this species has an expected life span of about 25yr (G.McPherson,
Northern Fisheries Centre, Cairns, pers. comm.).
Groupers are among the most important of the reef fish landings but little is known about the biology of species captured in the Pacific islands. The age, growth and
mortality of the coral trout Plectropomus leopardus on the central GBR has been studied by Ferreira & Russ (1992), while a detailed comparative study of the
population biology of P. leopardus from different locations on the GBR is given by Brown et al. (1994). Earlier estimates of the age and growth of P. leopardus were
made from lengthfrequency data (Goeden 1978) but this species like others on the GBR has otoliths with clear annual markings. Loubens (1978b, 1980) aged P.
leopardus from otolith annuli and included some ageatlength estimates for Epinephelus areolatus, E. fasciatus, E. hoedti, E. maculatus, E. merra and E.
tauvina. Little else has been documented on the biology of other groupers in the South Pacific. Morize & Caillart (1989) have made some preliminary investigations of
ageing juvenile and adult Epinephelus polyphekiadon (=microdon) from French Polynesia from otolith microstructure. Their results suggest that the periodicity of
primary growth increments is once every two days rather than daily and that the rate of increment formation is correlated with growth but not with fish length.
Longevities of groupers in the South Pacific based on the studies reviewed here appear to be typically in excess of 10yr.
The biology of other commonly exploited reef fishes in the South Pacific is more fragmentary with most examples confined to studies in Hawaii, the GBR and French
Polynesia. Like the groupers, jacks (Carangidae) are also a major component of reef fish catches but the biology of these species does not appear to have been the
focus of much attention from fisheries biologists in the region. The biology of the large jacks, Caranx melampygus and C. ignobilis, in Hawaii has been studied by
Sudekum (1984), who was able to age C. melampygus by primary growth increments in the otoliths. Also in Hawaii, a study was conducted on the movements,
distribution and growth rates of the goatfish Mulloidichthys flavolineatus by Holland et al. (1993) using tagging data. Other studies on reef fishes in Hawaii that have
generated life history data related to fisheries management include age and growth from otolith microstructure of the millet seed butterflyfish, Chaetodon miliaris
(Ralston 1976) and of the brick soldierfish Myripristis amaena, which Dee & Radtke (1989) suggested had a typical life span of 14yr. Growth and mortality of
another soldierfish Sargocentron microstoma in French Polynesia have been estimated from lengthfrequency data by AriasGonzalez et al. (1993).
Relatively few estimates of biomass or standing stocks have been made for reef fish populations in the Pacific islands and these are summarized in Table 9. The
majority of estimates are from the Australian GBR and the French territory of New Caledonia. The tech
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Table 9 Estimates of biomass or standing stocks of reef fish on South Pacific coral reefs.
Biomass (tkm−2)
Location Reef type Range mean Method Source
Woleai atoll (FSM) Atoll lagoon back reef 5–25 12 Depletion fishing Smith & Dalzell 1993
Australian GBR Inshore reef 92 Explosives Williams & Hatcher 1983
Australian GBR Midshelf reef 237 Explosives Williams & Hatcher 1983
Australian GBR Outer shelf 156 Explosives Williams & Hatcher 1983
Australian GBR Coral island fringing reef 17.5–195 97.5 UVC Goldman & Talbot 1976
Papua New Guinea (New Ireland) Fringing reef 43.5 Explosives A.Wright FFA. pers. comm.
Chesterfield Islands (Fig.2, 15) Coral islands and atolls 1.7–230 105 UVC and rotenone Kulbicki et al. 1990
New Caledonia Fringing reef 110 UVC Kulbicki 1988
New Caledonia Mid shelf reefs 78–96 90 UVC Kulbicki 1988
New Caledonia Barrier reef 92 UVC Kulbicki 1988
New Caledonia Ouvea Atoll 25–400 56.2 UVC Kulbicki et al 1994
Enewetak Atoll reef 42.5 UVC Odum & Odum 1955
Hawaii Fringing reefs 0.8–237 38.8 UVC Brock 1954
French Polynesia Fringing reefs 140 UVC Galzin 1987
Western Samoa Fringing reef slopes and lagoon 11.7–74.7 38.7 UVC Samoilys & Carlos 1991a
Fiji Barrier reefs 22.2–28.9 25.1 UVC Samoilys & Carlos 1992b
a,b. Biomass estimates of Serranidae, Acanthuridae, Scaridae, Lethrinidae & Lutjanidae only
UVC=underwater visual census, see text
niques used to estimate standing stocks include underwater visual census (UVC), controlled use of explosives, poisoning and shortterm intensive fishing experiments,
UVC techniques comprise underwater counts of reef fishes, either along a transect line of a given width or in a given radius from the observer, and estimation of lengths
of all individuals for subsequent conversion of numbers to weight through lengthweight equations. As a technique for estimating abundance and biomass of target
species, UVC has several limitations, as evident from the rarity of emperors in extensive counts around reefs of six different fishing grounds in Fiji (Jennings & Polunin
1995c), whereas these species constitute a major part of the reef catch. Explosive sampling is based on setting charges where fishes will be killed or stunned within a
known radius and then can be collected by divers following detonation. However, species such as eels, gobies and blennies in which the swimbladder is poorly
developed or absent are not as susceptible to this method of sampling. The use of poisons involves surrounding patch reefs or parts of a reef with nets then introducing
an ichthyocide such as rotenone on to the reef to kill all fish within the demarcated area. The objective of such intensive fishing over a short period of time is to reduce
the standing stock and hence the CPUE where factors such as growth mortality and recruitment are negligible. The reduction in
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CPUE can be correlated with cumulative catch (Leslie’s method) or fishing effort (DeLury’s method) to determine initial biomass or standing stock within the area being
subjected to intensive fishing (Ricker 1975).
From the examples in Table 9, the average standing stock biomass on Pacific reefs ranges from 12 to 237tkm−2. It is difficult to make any serious comparisons of
these figures, given the range of methods used and the restricted number of locations. The lowest mean figure from Woleai Atoll was obtained from a series of intensive
fishing experiments. Smith & Dalzell (1993) have suggested that their biomass estimates refer only to that fraction of the total biomass that was susceptible to the fishing
methods employed, namely spear fishing and drivein net fishing. Furthermore, the inner reefs of Woleai Atoll are fished regularly by the inhabitants of this island, who
rely on fish as a main source of protein and so the low biomass estimates may also reflect the removals by fishing. Smith & Dalzell noted that there was an apparent
correlation between biomass on the four reefs in their study and the time interval between the experiments and the last largescale community fishing events on these
reefs. The other examples in Table 9 are from reefs that are unexploited or were only lightly fished during the period of observations. The highest figures overall are
from the Australian GBR. Here the use of explosives, although not effective against certain species, permitted sampling throughout the water column and revealed the
large contribution from planktivores, especially fusiliers (Caesionidae), to the reef fish biomass. Based on the limited data in Table 9 it appears that unexploited or lightly
exploited coastal reefs may typically have standing stocks of reef fishes in the range 50 to 100tkm−2.
As the areas of reef, lagoon and coastal shelf in many locations in the South Pacific and elsewhere are readily obtainable from nautical charts, aerial photographs and
satellite images, it is often possible to express the catch from reefs in terms of production per unit area. This then gives an index of exploitation that can be compared
with other locations and from which it may be possible to obtain an indication of the sustainable yield. There are a number of examples from the South Pacific and
elsewhere in the tropics where the amount of fish and/or invertebrates taken from a given area of reef have been estimated. However, authors have used a variety of
techniques to estimate the fish and invertebrates harvested from a given area, and they have given different definitions of the area of reef being fished. Some workers
have restricted the definition of reef to include only actively growing hermatypic reef to depths ranging between 8 and 60m (see Russ 1991). Some authors have
estimated yields based on reef area and on area of shallow lagoon, which included tidal mangrove areas, seagrass beds and sand flats. Others have included the reef
and the adjacent shelf area to a depth of 200m in their estimation of reef yields. These boundary conditions clearly affect the results substantially. Using a depth limit of
40m for reef areas and 60m for other habitats, Jennings & Polunin (1995a) estimated yield ranges of 0.3–10.2tkm−2yr−1 (reef fish/reef area) and 0.2–3.4tkm−2yr−1
(reef fish/total area) in six Fijian fishinggrounds (qoliqoli).
The selection of species to be included in harvests creates particular problems when comparing reef yields between different locations. Catches by reef fishermen
may contain substantial catches of scombrid fishes such as tunas and mackerels (Table 3, p. 416) or snappers, groupers and other deep slope species caught away
from the reef, thus inflating reef yields. Reef yields may also include the shellfish and other invertebrates collected from the inshore reefs at low tide. This gleaning activity
can in some locations account for a significant fraction of the total harvest from a reef area, as in Western Samoa where invertebrates account for 36% of the total reef
landings (Wass 1982, Munro 1984), and in Fiji where they account for 72% by weight of artisanal catches and almost half of subsistence landings (Rawlinson et al.
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Table 10 Estimated yields from coral reef fisheries in the South Pacific.
Location Area fished Habitat (km2) Maximum depth Yield Source

type fished (m) (tkm−2yr−1)
Papua New Guinea Kavieng Fringing reefs and 207.7 30 0.42 Wright & Richards 1985
patch reefs
Papua New Guinea Port Moresby Barrier and fringing 116 40 5.0 Lock 1986b
reefs
Papua New Guinea Manus Fringing and patch 61.1 20 2.8 Chapau & Lokani 1986
reef
Papua New Guinea (total) All coral reefs 39940 30–60 0.21 Dalzell & Wright 1986
American Samoa Fringing reef 3.0 40 8.6–44.0 Wass 1982
Western Samoa Fringing reef 300.0 40 11.4 Zann et al. 1991
Tarawa (Kiribati) Atoll reef and lagoon 459.0 30 7.2 Mees et al. 1988
Ontong Java (Solomon Islands) Atoll reef and lagoon 122 na 0.6 BaylissSmith 1975
Nauru Fringing reef and reef 7.5 100 4.5 Dalzell & Debao 1994
slope
Niue Fringing reef and reef 6.2 60 9.3 Dalzell et al. 1993
slope
Vanuatu Fringing reef and reef 1063 100 1.9 David & Cillauren 1989
slope
Futuna Fringing reef 150 na 0.01 Based on data in Galzin 1985
Palau Fringing and barrier 450 20 1.7–3.0 Kitalong & Dalzell 1994
reefs
Fiji (Koro (41) & Lakeba (45)) Fringing reefs 8.4 na 5.0 BaylissSmith 1975
Fiji (Yanuca (116), Dravuni, Moala (62), Totoya Fringing reefs na na 0.3–10.2 Jennings & Polunin 1995a
(105), Nauluvatu (66))
Ifaluk Atoll (35) Atoll reefs and lagoon 5 na 5.1 Stevenson & Marshall 1974, based on
observations by Alkire 1965
1994). Despite these inconsistencies it is still possible to draw some conclusions from the estimates of yield from Pacific reefs summarized in Table 10. The observed
yields are primarily a function of fishing effort that is itself a function of population density. In the smaller Pacific island countries where marine produce has always been
a primary source of protein, population pressure may lead to relatively high levels of exploitation on nearshore stocks. Small islands such as Nauru and Niue, with
limited reef and shelf area and extensive fishing activity, have estimated yields of 4.8tkm−2 (Dalzell & Debao 1994) and 9.3tkm−2 (Dalzell et al. 1993) respectively from
the reef and shelf areas combined. The highest estimated annual fisheries yields were for fringing reefs in American Samoa, with a range of 8.6 to 44.0tkm−2 and a
mean of 27tkm−2.
Locations in Melanesia, where agricultural land is generally more abundant and population densities much lower, tend to have much lower yields from nearshore
fisheries. In PNG the annual total reef fisheries yield for the whole country was estimated to be 0.21tkm−2 (Dalzell & Wright 1986). Based on data presented by David
& Cillauren (1989), the total yield of reef fishes and invertebrates in Vanuatu amounted to only 0.16tkm−2. Higher annual yields have been recorded at individual
locations in PNG such as Manus (Fig. 2, 54) (≈3.0tkm−2) (Chapau & Lokani 1986) and at the capital city Port Moresby (≈5.0tkm2) (Lock 1986c), but these are still
relatively modest when compared with islands in Polynesia and Micronesia. The limited data suggest that finfish yields in the range 5 to 20tkm−2yr−1 are probably
sustain
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able in the long term. If coastal reefs typically have standing stocks in the range 50 to 100tkm2, then annual sustainable harvests are likely to represent between 5 and
40% of the standing stock (fishable teleost biomass), although we acknowledge that this amounts to little more than speculation given the limits of the present data on
standing stocks and yields.
A log book study of six Fijian fishing grounds (qoliqoli) established that at least up to 10.2tkm−2yr−1 (reef fish/reef area up to 40m depth), yield was linearly related
to a rescaled index of effort (Jennings & Polunin 1995b), indicating that such harvests are sustainable and indeed they appear to have been maintained for some years in
the most fished area (Jennings & Polunin 1995a). Although catch composition varied between grounds, there was no evidence that lessfavoured species were
increasing at the more intensely fished sites (Jennings & Polunin 1995a).
The most comprehensive study of fisheries resources and potential MSY in the region has been conducted by the French scientific organization ORSTOM in Ouvea
Island (Fig. 2, 75) (Kulbicki et al. 1994), one of the Loyalty Islands archipelago that lie to the east of New Caledonia. From a series of underwater observations and
experimental fishing surveys, Kulbicki et al. (1994) estimated that the total standing stock of reef and lagoon fish was 46500t of which 12500t was found on the reefs.
Kulbicki et al. (1994) estimated that the MSY for the fishable stock might be as high as 4300tyr−1 or about 10% of the total standing stock, but recommended that a
more conservative yield of 1000tyr−1 would be a better management objective. This would amount to a relative yield of about 1.1tkm−2yr−1 from the lagoon and reef
area combined. The present volume of landed catch from Ouvea is only 50tyr−1 or onetwentieth of the predicted MSY.
Other attempts to estimate MSY for reef fisheries have used modifications of simple surplus production models, either the simple parabolic form initially developed
by Schaefer (1954) or the exponential modification of Schaefer’s model proposed by Fox (1970). Both forms of the model are based on the assumption that growth in
fish populations conforms to a logistic or Sshaped curve, with maximum production from a given stock at some intermediate population density (Ricker 1975). The
sustainable yield is the level of fishing effort that takes only the annual surplus production from the population without depleting the biomass. In the Schaefer model the
relationship between catch and fishing effort conforms to a symmetrical parabola with MSY when the population is reduced through fishing to half of the unexploited
biomass. The exponential Fox model is an asymmetrical dome shaped curve where MSY occurs when the population is reduced to about 40% of the unexploited
biomass.
Conventional versions of the Schaefer and Fox surplus production models require time series of catch and effort data to generate MSY and optimum fishing effort.
While long time series of reef fish catch data (15–25yr) are available for countries such as Palau, French Polynesia and Fiji, there are no accompanying data on fishing
effort due to the multiplicity of gears involved in the fishery. A shorter, incomplete set of catch and fishing effort data for Tarawa Atoll (Kiribati) was analyzed by
Yeeting & Wright (1989). A direct measure of fishing effort was not available but an index of effort could be expressed as number of canoes or skiffs, and numbers of
gears such as handlines and gill nets. Catch was broadly separated into reef catch, lagoon catch and ocean catch. Ocean catches were predominantly tunas caught by
trolling and, not surprisingly, mean annual catch rates showed no relationship to annual fishing effort. Mean annual catch rates from the atoll’s lagoon and reef stocks
showed an inverse linear relationship with effort in gear numbers and in terms of vessel numbers, and catch and effort data could be fitted with Schaefer curves. The
conclusions from this analysis were that the MSY of reef and lagoon fish stocks in Tarawa Lagoon was about 2800tyr−1 or a yield from the total fished area (141.8km2
reef and 330.4km2 lagoon) of 6tkm−2yr−1.
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Lock (1986bc) fitted a Fox surplus production model using spatial variation in fishing effort and catch on a number of different reefs in the Port MoresbyDaugo
Island reef fishery over a 1yr period, to compensate for the lack of a time series of catch and fishing effort data. This method was adopted from Munro (1983) who
developed this technique to estimate the MSY for the nearshore canoe fishery in Jamaica. Lock’s estimate of MSY for the Port MoresbyDaugo Island Fishery was
524tyr−1 or a yield of 7.6tkm−2yr−1. Munro (1984) also used this approach with data from Wass (1982) to fit a surplus production model to yield data for finfish and
invertebrates from reefs around Tutuila Island in American Samoa. Although total harvests from Tutuila (Fig. 2, 108) are not known, Munro concluded that the data
indicated an annual finfish MSY of 20tkm−2yr and an invertebrate yield of 15 tkm−2yr−1.
Another approach to investigating fisheries yield from coral reefs has been to construct a trophodynamic model of a reef system. Polovina (1984a) developed an
ecosystem box model, ECOPATH, to estimate the biomass budget of an ecosystem given inputs that specify the ecosystem components, together with their mortality,
diet and energetics value. This approach was used to model a coral reef ecosystem at French Frigate Shoals in the Northwest Hawaiian Islands (Polovina 1984b,
Atkinson & Grigg 1984, Grigg et al. 1984), and suggested that fisheries yield might be maximized by harvesting low in the food chain, particularly if top carnivores can
be cropped to release predator pressure on selected prey. One use of trophodynamic modelling of reef communities has been the indication that fishes around reefs
contain substantial proportions of the total amount of inorganic nutrient elements such as nitrogen stored within biomass (Polunin 1996). The implication is that in some
cases intense fishing may lead to nutrientdepletion because of the removal of a major reservoir within the reef system, recovery of which may be slow. The extent to
which reef fish productivity may depend on nutrient inputs, however, is unknown. The opportunity to examine changes in biomass and turnover of reef fish stocks in
nutrientpoor and upwellingenriched regions offered by the Pacific has apparently not been explored. The potential role of planktonic inputs in the production of
groupers, snappers and other important reef predators is being explored, however, by using carbon, nitrogen and sulphur stable isotopes (N.V.C.Polunin unpubl. data).
Socioeconomic developments
Shallow water reef fisheries, like most coastal fisheries in the Pacific, remain the preserve of smallscale artisanal fishermen. Even in the commercial sectors of most
countries, the gears employed are largely nonmechanized, and in the Pacific, most reef fish catches are generated from handlines, spears, gill nets or drivein nets. The
only exception has been the proliferation of fish corrals in the Tuamotu Archipelago of French Polynesia, where these gears are used to catch large volumes of reef
fishes to satisfy the demand for fish on Tahiti, the principal island in the territory. Elsewhere, fishermen have sought to improve catches through the greater use of
modern fishing gears such as monofilament lines and nets, diving equipment and more reliance on outboard motors to range over wider areas. Governments have sought
to encourage greater fisheries production from reef and other coastal fisheries by providing better facilities for fishermen to dispose of catches and in some instances
provision of easy credit or “soft” loans to buy vessels and equipment.
In the past, most catches from nearshore reef fisheries were consumed at or close to the landing site but, as indicated earlier, there was a greater dispersal of landings
from their point of origin to other domestic and international markets as national economies developed and urbanisation increased. Besides the examples given earlier
for French Polynesia, Cook
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Islands and Western Samoa, reef fishes are now exported from Kiribati to Hawaii and Nauru, from Solomon Islands to Japan, and from Fiji to Tonga and New
Zealand. A trend in the western part of the region is the development of live reef fish fisheries for the restaurant trade in Southeast Asia. Reef fishes, such as coral trout,
other groupers and Napoleon wrasse, are caught around the coast of PNG and shipped live by air and sea to markets in Hong Kong, mainly for the restaurant trade,
although there is also a demand for live stonefish (Synanceia verrucosa and S. horrida) for traditional Asian medicines as well as for food (Richards 1993). A similar
type of operation was conducted from Palau to Hong Kong before concern over levels of fishing pressure forced the closure of the fishery (Johannes 1991). Interest
has also been shown in developing live reef fish exports from Tuvalu (I.Keay, Tuvalu Fisheries Division, pers. comm.) and from the Australian GBR (A.H. Richards,
Forum Fisheries Agency, pers. comm.) and Richards (1993) has suggested that as reef fish resources are increasingly depleted in the South China Sea region,
companies based in Taiwan, Hong Kong and Singapore will venture increasingly into the South Pacific in search of groupers and Napoleon wrasse.
There is also growing interest in the Pacific islands in developing export fisheries for the international aquarium fish trade. Aquarium fish fisheries are operating in Fiji,
the Cook Islands, Vanuatu, Palau, Kiribati, Tonga and the Marshall Islands. The common target species of aquarium fish fisheries are the smaller reef species not
normally targeted heavily for food. Pyle (1993) lists 10 families (Pomacanthidae, Chaetodontidae, Acanthuridae, Labridae, Serranidae, Pomacentridae, Balistidae,
Cirrhitidae, Gobiidae and Blenniidae) as the most important to the aquarium trade. Over 240 species of reef fish are listed in the export figures reported from Palau
(Anon 1993b), with the most common being the damselfishes (Pomacentridae), Chrysiptera cyanea, Chromis albipectoralis and Dascyllus aruanus, which are of
little unit value, price range US$0.2–0.3 per fish. However, this is much higher than the price received for reef fishes sold for food and comparatively rare specimens
such as Ctenochaetus tomiensis may be worth as much as US$25per fish.
Improvements in fishing power of gear and growth of human populations have in many locations been paralleled by declines in stocks, catch rates and, in some
cases, landed volume of reef fishes. Certain species may be extremely vulnerable to particular fishing gears. As stated earlier, the combination of spear fishing and
SCUBA gear is believed to responsible for the extinction of Bolbometopon muricatum and Cheilinus undulatus at Guam and the largescale population decline of
these species on Palau. Other cases of finfish stock depletion have been reported in the Pacific. These include declines in reef and lagoon fish stocks in Palau (Johannes
1981, 1991), reef fishes and lagoon fishes in Kiribati (Yeeting & Wright 1989), bonefishes, milkfishes and parrotfishes in the Cook Islands (Anon. 1988a, J.
Dashwood, SPC Fisheries Programme, pers. comm.), various grouper stocks in French Polynesia (Bell 1980), Tokelau (Hooper 1985) and the Cook Islands (Sims
1990), and reef and small pelagic fishes in Western Samoa (Helm 1992).
In many locations in the Pacific, exploitation of reef resources is regulated by communities, particularly via traditional concepts of marine tenure (see contributions in
Ruddle & Johannes 1985 and South et al. 1994). In some locations, customary ownership of fishery resources has almost disappeared, resulting in open access to
nearshore reef resources. Fisheries management and development must account for traditional ownership where it exists, and this may be formalized in legislation as in
Fiji (Adams 1993c, Adams 1996). Contemporary approaches to managing reef fisheries in some locations in the region may include very detailed fisheries regulations
and ordinances that specify closed seasons, areas closed to fishing, size limits for certain species and mesh size limits for fishing gear. Such approaches are
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likely to be more enforceable in the commercial sector than at the community or village level.
Apparently, the only objective attempt to assess the robustness of traditional management in the face of resource pressure has been that of A.Cooke and
N.V.C.Polunin (unpubl; see also Cooke (1994) and Cooke & Moce (1995)). An index of “management aptitude” was derived from the responses of eight Fijian
qoliqoli managers to a set of specific questions including queries as to their use of information for management, approach to goodwill payments, work with the Fisheries
Division and management and enforcement measures taken. Of four qoliqoli subject to high commercial fishing pressure, two showed high aptitude and two low
aptitude. Among the qoliqoli studied, those with highest aptitude showed, in particular, evidence for liaison and collaboration with the Fisheries Division. The inference
was that those exhibiting low aptitude might benefit from some form of comanagement with the Fisheries Division.
Coastal reef finfish catches will continue to be the main source of subsistence protein for most Pacific island countries for the foreseeable future, but there is likely to
be an increasing volume of high value species being transported to domestic urban and tourist centres and exported to overseas markets. Problems are likely to occur
in countries where yields from coral reef fisheries cannot keep pace with population growth and where there are no major developments in targeting offshore fish stocks
or aquaculture. Those islands most at risk from this “Malthusian over fishing” (as defined by Pauly 1990) of reef fish will be those with a high human population in
comparison with the available reef and lagoon area. There is not yet any comprehensive quantification of coastal fisheries habitat areas for the Pacific islands, so this
ranking cannot yet be made, but islands such as Saipan, Upolu, Rarotonga and Tarawa, for example, would definitely appear to be in a highrisk category.
Deepslope fisheries
Description
Beyond shallow reef slopes, in depths where hermatypic corals do not flourish, lie the deep reef slopes. These are usually areas of sand and coral boulders, but with
other sediments also present depending on island type and proximity to rivers and alluvial deposition. The deep reef slope typically starts at about 80m and extends to
about 400m depending on the steepness of the island shelf. The fish community of the deep reef slope is simpler than the neighbouring shallow reef and comprises
mainly large carnivorous species of snappers (Lutjanidae), groupers (Serranidae), emperors (Lethrinidae) and jacks (Carangidae). Catches of snappers from the deep
reef slope are dominated by members of the genera Pristipomoides and Etelis (referred to here collectively as eteline snappers) that are high quality fish with a high
demand from overseas markets in Japan and Hawaii. These deepslope fish stocks have, until recently, been lightly exploited or unexploited throughout most of the
South Pacific region. Exceptions are the limited subsistence fisheries for deepslope snappers and oil fish (Ruvettus pretiosus) at some Polynesian atolls (Wankowski
1979) and in Hawaii where deepslope fish stocks have been continuously exploited for over 50yr (Ralston & Polovina 1982, Polovina 1987, Shomura 1987).
During the early 1970s, the South Pacific Commission commenced surveying the deep reef slope stocks of the Pacific islands and demonstrating techniques useful
for exploiting these stocks (Dalzell & Preston 1992). Initial surveys in Polynesia and Melanesia revealed the existence of fishable stocks but the techniques used were
not appropriate for the Pacific
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islands. From 1979 onwards the Commission propagated the use of more appropriate fishing technology, based around small dieselpowered dories and manual fishing
reels first developed in Western Samoa (Gulbrandsen 1977). The efforts of the Commission and its extension programme led to the establishment of deepslope
fisheries in many locations in the South Pacific, including Tonga, Fiji, Vanuatu, American Samoa, Western Samoa, Solomon Islands, PNG, Federated States of
Micronesia and French Polynesia.
Not all of these fisheries have persisted and the reasons why they have failed in various locations are a mixture of both stock depletion and socioeconomic factors.
The most successful deepslope fishery is probably the fishery in Tonga, which began on the nearshore slopes of the archipelago but graduated to the numerous
seamounts in the Tongan EEZ from which most of the Tongan deepslope catch now originates. The deepslope fishing fleet is composed of wooden dories between 6
and 11m in length, powered by 20hp Yanmar diesel engines. The boats were constructed locally with credit supplied from two United Nations’ agencies. All boats use
between four and five wooden handreels, most commonly baited with the commercial longline bait of saury (Cololabis saira).
Descriptions of the Tongan fishery are given by Langi & Langi (1987) and Latu & Tulua (1991). The dories make voyages lasting 5days, of which 2days are
traveling time and three days are spent fishing on seamounts. On average between three and four reels are deployed during a fishing trip for a period of about 7h per
day. Each vessel completes about 30 fishing trips per year. Most of the dories are based in Tongatapu, with a small number of boats landing fish into Vava’u. The
fishermen sell their catch to fish buyers, some of whom also own fishing vessels. About twothirds of the catch is sold locally, while the prime species Etelis coruscans
and Pristopomoides filamentosus are air freighted to markets in Hawaii. Initially, about 40 dories were operating in the fishery, but the fleet has now shrunk to
between 15 and 20 vessels: some boats have been used for other purposes such as bêchedemer collection or simply were no longer seaworthy. Annual landings from
the Tongan deepslope fishery amount to between 210 and 514tyr−1 with current production at around 250tyr−1.
The deepslope fishery in Fiji has not been as well documented as the Tongan fishery but was nearly as large during the mid 1980s in terms of production volume.
Furthermore, the Fiji fishery was innovative in the use of larger more sophisticated fishing vessels, using commercial fishfinding sonars and deploying bottom longlines
and hydraulically operated reels rather than the simple wooden handreel. The Fijian fishery was also the first in the South Pacific to explore the possibility of exporting
fish to more lucrative overseas markets. Lewis et al. (1988b) provide the best description of the Fiji deepslope fishery prior to its demise after 1987. The Fiji deep
slope fleet comprised one 20m Hawaiian longliner, four larger local vessels (three dropline and one longline) and a number of 9m dories, similar to those used in
Tonga. The larger vessels deploying hydraulic reels used lines with five or more hooks per line. Bait used throughout the fishery was skipjack rejected by the local
cannery. The larger vessels used a palu, or chum bag, to aggregate fish and increase catch rates. At the peak of the Fiji fishery about 200t of deepslope fishes were
landed annually, with about 75% of this sold overseas. Disruption in airline scheduling following political events in 1987 was a serious setback to the fishery, where
profit margins were not large. However, the vessels involved in the fishery began to shift from demersal fishing to pelagic longlining to catch large highvalue tunas such
as yellowfin (Thunnus albacares) and bigeye (T. obesus). These species can be caught more reliably than deepslope fishes, realize a much better return on overseas
markets and stocks are not nearly as limited as stocks on the deep slope. The expansion of the deepslope fishery in Fiji was based largely on catches from unexploited
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stocks, where catch rates could fall by one order of magnitude in a short period of time, particularly when fishing on seamounts.
The Vanuatu deepslope fishery was based on the Village Fisheries Development Plan (VFDP) that was conceived as a strategy to increase the supply of fresh fish
from village based fisheries in the country. As part of the overall plan, locally built wooden dinghies and catamarans were equipped with the Samoan handreel and
various incentives were offered to village groups to become involved in the fishery, such as dutyfree gasoline. In recognition of the fact that Vanuatu villagers were not
used to fishing outside the shallow reef zone, a training centre was established in Luganville (Fig. 2, 125), to impart the skills required to become an artisanal fishermen
targeting deep slope fishes. The fishery expanded from just six boats in 1982 to 180 by 1988, although not all boats were engaged in fulltime fishing. Descriptions of
the deepslope fishery in Vanuatu are given by Schaan et al. (1987) and Carlot & Nguyen (1989). The volume of landings in the Vanuatu fishery between 1982 and
1988 ranged from 10 to 86tyr−1 with a mean of 50tyr−1, while a survey of commercial fisheries in Vanuatu during 1992 suggested an annual production from the deep
slope fishery of about 80t (Anon. 1992b). Dalzell (1992a) noted that in response to declines in catch of deepslope fishes, Vanuatu fishermen were now targeting
shallow reef species. Although the VFDP was planned to increase fish supply for the village population, much of the deepslope catch is now sold to the restaurant and
hotel trade in Port Vila (Fig. 2, 81) and Luganville.
PNG has by far the largest resource of deepslope species in the South Pacific but only one commercial fishery, on the north coast of the mainland, was ever
successfully established there. The fishery landed between 5 and 20tyr−1 between 1983 and 1985, before going into decline after government support for the fishery
was reduced (Chapau & Dalzell 1991). A combination of factors contributed to the decline of the American Samoan fishery, which was comparable in scale to the
fishery in northern PNG, but where landings were exported to Hawaii to realize greater profits. Among the factors responsible for the decline in the fishery were a fall in
catch rates of deepslope stocks, volatility of the prices on the Hawaiian market, delays in payment for export catches and competition from purseseine bycatch on
the domestic market (Itano 1991).
Elsewhere in the South Pacific, small amounts of deepslope fishes are caught for local markets. Landings of deepslope fishes in French Polynesia are mainly from
recreational fishermen and landings range from 0.5 to 10t annually (Wrobel 1988). Between 40 and 80t of deepslope fishes are landed annually in New Caledonia for
domestic consumption (Anon 1994a). Recently, a commercial survey of seamounts and banks in the Tuvalu EEZ has led to catches of deepslope fishes, some of
which have been marketed in Hawaii (Anon 1993c). Small amounts of deepslope fishes have been caught and airfreighted to Japan and Hawaii from the Federated
States of Micronesia, but production has not been consistent (P.Dalzell unpub. data).
Catch composition
By contrast to shallow reef fish fisheries, which have been largely ignored by fisheries scientists in the South Pacific, deepslope species have been the focus of a
considerable amount of research and monitoring. Dalzell & Preston (1992) present the most coherent data set on composition of deepslope fishery catches throughout
the Pacific. These data (Table 11) are based on surveys conducted by the South Pacific Commission on what are essentially unexploited stocks. Nearly all the Pacific
islands are included in this data set, notable excep
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tions being Guam, Nauru and Pitcairn. The snappers, or Lutjanidae, are divided into two groups, following the taxonomy proposed by Johnson (1980); the subfamilies
Etelinae and Apsilinae, or deepslope snappers, and the subfamilies Lutjaninae and Paradichthyinae, or shallow water snappers.
The Lutjaninae are more a feature of shallow lagoon habitats but species such as Lutjanus bohar and L. argentimaculatus migrate down the deep reef slope as
they increase in size (Wright et al. 1986). Indeed, of the total number of fishes caught in the Commission surveys, L. bohar was the commonest species (4.6%),
followed by Caranx lugubris, Pristipomoides filamentosus, Etelis carbunculus and Pristipomoides flavipinnis that together formed over 20% of the total catch.
A further eight snapper species (Pristipomoides zonatus, P. multidens, P. auricilla, Etelis radiosus, E. coruscans, Lutjanus gibbus and Aphareus rutilans),
formed a further 20% of the total.
Factors such as position or longitude, average depth fished, seasonally, island type and size and fishing intensity will have an effect on the composition of catches.
Dalzell & Preston (1992) compared species composition between catches at high islands and catches around atolls. They showed that catches from high islands contain
a significantly greater amount of the commercially valuable eteline snappers than at atoll sites. Furthermore, catches at atoll sites contained greater numbers of sharks,
which have little or no commercial value. Regional differences also exist between the species composition of deepslope catches from the Pacific islands. We have
summarized data on catches from the South Pacific Commis
Table 11 Percent composition of deepslope catches from different locations in the South Pacific.
Country/Territory Etelinae Lutjaninae Lethrinidae Serranidae Carangidae/Scombridae Gempylidae Sphyraenidae Other teleosts Sharks
American Samoa 42.4 18.1 14.9 2.1 12.4 0.7 8.9 0.8 0.0
Belau 36.0 11.8 4.1 12.4 15.2 7.0 1.7 3.5 5.2
Cook Islands 50.7 2.0 1.2 9.4 9.9 4.7 0.1 6.7 15.2
Federated States of Micronesia 21.7 18.6 4.4 7.4 18.5 2.4 1.0 4.9 21.2
Fiji 24.6 11.9 5.4 8.1 15.1 2.7 5.2 1.5 25.6
French Polynesia 28.7 2.5 0.2 30.2 19.7 14.1 0.1 1.6 3.0
Kiribati 13.5 32.8 3.4 21.6 5.5 5.4 0.4 0.7 17.1
Marshall Islands 8.5 14.3 6.5 10.1 8.1 1.1 0.3 2.3 48.9
New Caledonia 24.4 9.6 19.4 11.4 5.6 0.0 2.8 0.2 25.3
Niue 10.1 27.2 11.2 13.5 9.2 3.9 1.2 15.3 3.3
Northern Marianas 60.3 0.0 0.1 0.5 34.4 0.0 0.0 0.0 0.0
Papua New Guinea 49.2 16.1 3.8 7.2 7.0 0.5 0.9 0.4 15.3
Solomon Islands 61.0 14.3 0.4 10.8 1.9 0.0 7.8 3.8 0.0
Tokelau 21.8 2.9 6.1 6.9 27.9 5.5 1.5 2.3 25.4
Tonga 49.0 3.9 20.2 13.8 5.5 0.8 1.1 0.5 5.0
Tuvalu 17.1 10.2 0.9 10.1 12.6 32.8 0.7 0.5 15.2
Vanuatu 45.5 10.6 2.1 19.1 3.9 4.6 0.2 1.3 12.9
Wallis & Futuna 56.8 4.6 5.8 12.2 11.6 0.0 0.1 0.3 8.8
Western Samoa 45.9 7.1 1.2 5.6 5.0 25.0 0.0 3.8 6.5
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Table 12 Composition by species of three families taken by deepslope handline fishing in the South Pacific region.
Percentage composition by subregion

Family and species Melanesia Micronesia Polynesia
LUTJANIDAE
ETELINAE/APSILINAE
Aphareus furca 0.05 0.82 0.90
A. rutilans 7.98 6.27 8.34
Aprion virescens 1.49 4.41 6.49
Etelis carbunculus 6.76 20.56 9.47
E. coruscans 4.71 15.02 5.53
E. radiosus 2.79 0.20 0.89
Pristipomoides kuskarii 3.13 2.91 0.00
P. amoenus 1.20 0.52 0.21
P. auricilla 1.59 5.82 27.63
P. filamentosus 23.41 10.83 10.84
P. flavipinnis 18.61 9.32 4.80
P. multidens 22.99 1.11 0.98
P. zonatus 2.80 17.29 18.19
Others 2.50 4.92 5.73
LUTJANINAE
Lutjanus argentimaculatus 8.76 18.08 2.65
L. bohar 22.59 43.38 33.18
L. gibbus 11.85 29.66 15.20
L. kasmira 2.16 1.85 32.06
L. malabaricus 18.86 0.25 0.22
L. monostigma 8.64 0.95 0.49
L. rufolineatus 2.04 4.38 1.57
Others 25.10 1.44 14.62
LETHRINIDAE
Gymnocranius japonicus 5.23 0.30 0.82
Lethrinus chrysostomus 11.80 0.89 53.18
L. kallopterus 0.81 5.33 4.00
L. miniatus 15.90 32.99 17.01
L. reticulatus 0.15 3.11 0.00
L. variegatus 1.76 11.09 5.94
Wattsia mossambica 52.09 27.51 7.99
Others 12.27 18.79 11.07
SERRANIDAE
Cephalopholis aurantia 0.25 1.25 2.32
Epinephelus areolatus 4.39 8.26 4.01
E. chlorostigma 7.05 2.35 0.23
E. cometae 8.13 0.16 0.41
E. fasciatus 0.00 0.55 1.82
E. flavocaeruleus 1.53 1.76 0.27
E. magniscutis 0.49 0.63 0.00
E. miliaris 3.79 18.23 9.43
E. morrhua 27.35 14.44 12.48
E. retouti 1.13 0.47 10.02
E. septemfaciatus 4.73 0.08 5.15
Saloptia powelli 0.89 0.16 5.60
Variola louti 7.24 8.10 11.75
Others 33.02 43.58 36.49
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sion’s surveys presented by Dalzell & Preston to highlight differences between four common components of deep slope catches, namely eteline snappers, lutjanine
snappers, emperors and groupers, with respect to the three archipelagic groupings of Melanesia, Micronesia and Polynesia (Table 12). This effectively splits the Pacific
into the small high islands and atolls north of the equator (Micronesia), the large high islands in the west of the Pacific and south of the equator (Melanesia) and the small
high islands and atolls of the central Pacific that lie to the south of the equator (Polynesia)
In the small high islands and atolls of Micronesia, north of the equator, deepslope catches of eteline snappers are dominated by Pristipomoides auricilla and P.
zonatus. In similar habitats in the central and southern Pacific P. zonatus is also a major component in deepslope catches along with Etelis carbunculus and
Pristipomoides filamentosus. Catches on the slopes of the large Melanesian islands are dominated by P. multidens and to a lesser extent by P. filamentosus. As
with the etelines, there are subregional differences in the lutjanine snapper composition from deepslope handlining. In the Polynesian islands the small blueline
snapper, Lutjanus kasmira, and the red bass, L. bohar, each form about onethird of the lutjanine snapper catch, with the other major contribution being from L.
gibbus. In similar habitats in Micronesia, L. bohar is the dominant lutjanine snapper, along with L. gibbus and L. argentimaculatus. On the larger slope areas of the
Melanesian islands, L. bohar, L. argentimaculatus and L. gibbus are still dominant features of the snapper catch but dominance is shared with L. malabaricus.
Over half of the catch of emperors from the deep slopes around the large Melanesian islands comprises the deepslope bream, Wattsia mossambica. Other
dominant emperors included large and readily identifiable Lethrinus olivaceus (= miniatus) and L. miniatus (= chrysostomus). L. olivaceus and Wattsia
mossambica were also dominant features of the emperor catch from Micronesian islands along with Lethrinus kallopterus and L. variegatus. The dominant feature in
the catch from Polynesian islands was L. miniatus followed by L. olivaceus. Wattsia mossambica formed only a small portion of the lethrinid catches around
Polynesian islands.
The proliferation of species in the family Serranidae and the difficulties in identifying species, particularly in the field, are reflected in the relatively large percentages in
the other species category (Table 12) . However, grouper catches from the Pacific islands tend to be formed mainly from the following species: Epinephelus morrhua,
E. miliaris, E. retouti and Variola louti. In Melanesia, Epinephelus morrhua was clearly the dominant grouper species in deepslope catches, while in Micronesia
dominance was shared between E. miliaris and E. morrhua, and in Polynesia, among E. morrhua, E. miliaris and E. retouti. The coral cod Variola louti was
common to grouper catches in all three locations, while Epinephelus cometae and E. chlorostigma were dominant features of catches in Melanesian waters, and E.
areolatus of catches from Micronesia. Another feature of deepslope catches from Melanesian and Polynesian islands, particularly from unexploited fishing grounds is
the giant grouper E. septemfasciatus. This species is among the first to be fished out in deepslope fisheries where effort is particularly heavy.
Brouard & Grandperrin (1985) classified catches of deepslope catches in Vanuatu according to depth. Shallow species were defined as those in waters < 120m
and included many species commonly found on coral reefs such as squirrelfishes, small groupers, emperors and snappers. Species in waters of intermediate depths
(120–240m) consisted mainly of eteline snappers of the genera Pristipomoides and Paracaesio, and the larger emperors, lutjanine snappers and groupers. Deep
water species (>240m) included the three Etelis species, large groupers such as Epinephelus septemfasciatus, hexanchid sharks and oil fish
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(Ruvettus pretiosus). Sundberg & Richards (1984) arranged the 10 most common species in deepslope (80–300m) catches from northern PNG from common to
less common as follows: Lutjanus bohar, Lethrinus miniatus, Lutjanus erythropterus, Aphareus rutilans, Lutjanus malabaricus, L. argentimaculatus, Wattsia
mossambica, Pristipomoides multidens, Etelis coruscans and E. carbunculus.
Catch rates
The most comprehensive summary of catch rates from fishing on Pacific island deepslope stocks is given in Dalzell & Preston (1992). It should be noted, however,
that these catch rates are for mainly unexploited stocks and do not apply to commercial fishing. Catch rates ranged from 0.5–19.0kgline−1h−1 with a mean of
7.0kgline−1h−1. These gross catch rates include sharks, which are sometimes discarded and not recorded in catches. Catch rates for teleost fish only ranged from 0.3–
14.5kgline−1h−1, with a mean of 6.6kgline−1h−1. Average catch rates around high islands and atolls were 5.5 and 6.8 kgline−1h−1 respectively but there were no
significant differences between these means.
Average catch rates in commercial fisheries and survey fisheries in the South Pacific are also included in Table 5 (p. 418). Catch rates in the Tonga deep slope
fishery ranged from 2.76–13.3kgline−1h−1 with an overall mean of 6.4kgline−1h−1 for the years 1986–92 (Mees 1994). Sustained fishing on deepslope stocks over a
period of 1 yr in northern PNG by Richards & Sundberg (1984) produced catch rates ranging from 0.6 to 11.24kgline−1h−1, with a mean of 3.1kgline−1h−1. Similarly,
Chapau (1988) reported an average catch rate of 3.7kgline−1h−1 in the small commercial deepslope fishery based near Wewak (Fig. 2, 114) in northern PNG
between 1983 and 1985. A similar smallscale operation fishing around Pohnpei (Federated States of Micronesia) and the nearby atolls of Ant and Pakin (Fig. 2, 2)
between 1983 and 1986 produced catch rates that ranged from 3.9 to 5.5kgline−1h−1 with a mean of 4.7kgline−1h−1 (McCoy 1990). A smallscale pilot commercial
fishery operating around Pohnpei in 1989 experienced catch rates of 0.7–5.1kgline−1h−1 with a mean of 3.0kgline−1h−1 (P.Dalzell unpub. data). Survey fishing of banks
and seamounts around Tuvalu during 1992 and 1993 experienced catch rates ranging between 1.72 and 9.62kgline−1h−1 with an overall mean of 6.1kgline−1h−1
(Anon. 1993c). Catch rates experienced by village fishermen fishing on deepslope stocks in Vanuatu in the mid 1980s ranged from 0 to 5.00kg line−1h−1 with a mean
of 1.44kgline−1h−1. Average monthly catch rates for handline fishing on the Chuuk Outer Banks (Fig. 2, 17) ranged from 1.3–4.57kgline−1h−1 with a mean of
2.30kgline−1h−1. Catches comprised both deepslope and shallow water reef fishes. Sustained commercial fishing on the deepslope stocks of Niue between 1988 and
1990 generated average monthly catch rates ranging between 2.1 and 8.5kgline−1h−1, with an overall mean of 5.5kgline−1h−1.
Less information is available on catch rates from longline fishing on deepslope stocks and most of the fishing refers to surveys of virgin stocks. The limited
information available has been summarised in Table 13. Average catch rates ranged from a low of 6.8kg 100 hooks−1 on the Chuuk Outer Banks to a high of 124 kg
100 hooks−1 on the shelf area off the south coast of Espiritu Santo (Fig. 2, 22) in Vanuatu. The catch data from Fiji are from a commercial fishing operation that fished
on the outer banks and sea mounts in Fiji’s EEZ and are probably more indicative of the returns from this method of fishing on deepslope fish. Also included in Table
13 is a summary of longline fishing in the lagoon of New Caledonia in waters between 5 and 40m in depth. Although in shallow waters the composition of the New
Caledonia target stocks at the family level was closer to that from deepslope fishing
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Table 13 Summary of catch rate and catch composition data for longline fishing on coral reef and associated stocks in the South Pacific region.
CPUE (kg 100 hooks−1) Principal catch components

Location Target stock range mean Source
New Caledonia Shallow reef species 3.0–12.2 8.2 Lethrinidae, Kulbicki &
Serranidae, Grandperrin 1988
Labridae,
Lutjanidae
Chuuk Outer Banks Deep slope species 1.6–12.3 6.8 Lutjanidae, Diplock & Dalzell 1991
Carangidae,
Lethrinidae,
Tonga Deep slope species na 11 Lutjanidae, Mead 1987
Serranidae,
Carangidae,
Sharks
Fiji Deep slope species 12.5–29.2 19.1 Lutjanidae, Walton pers. comm.
Serranidae
Vanuatu (Paama (Fig 2, 76) & Espiritu Santo) Deep slope species 10–92.5 33.7 Lutjanidae, Serranidae, Fusimalohi & Preston 1983
Sharks
Vanuatu (Espiritu Santo) Deep slope species 15–389 124.6 Lutjanidae, Serranidae, SPC unpub. data
Lethrinidae,
Carangidae
than from reef areas, their inclusion in Table 13 allows for some comparisons to be made with deepslope catch rates.
Seasonal trends in catch rates of deepslope fishes have been reported from Vanuatu (Brouard & Grandperrin 1985) and from Tonga (Latu & Tulua 1991). In
Vanuatu catch rates for E. carbunculus and E. coruscans reached a maximum between February and June, with minima towards the year’s end. Maximum catch
rates of Pristipomoides flavipinnis and P. multidens were observed between April and July. By contrast the CPUE for Lutjanus malabaricus was at a maximum
during December and lowest in June and July. There was little evidence of seasonality in CPUE from the total catch from the deepslope fishery in Tonga; however,
individual species had very clear seasonal maxima in Tonga. The CPUE of both Pristipomoides filamentosus and P. flavipinnis were lowest in September and
highest in DecemberJanuary. Etelis coruscans was observed to have two peaks in CPUE, during May and during November, while only a single peak in CPUE was
observed for E. carbunculus during November. Two seasonal peaks in CPUE were also observed from catches of Lethrinus chrysostomus, during February and
July. The seasonal pattern for Epinephelus morrhua resembled that of the Pristipomoides spp., but with a peak in April and a low point in August. The seasonal
pattern for Epinephelus septemfasciatus showed a strong peak in CPUE during May.
Variation in catch rate with depth and time of day has been investigated in Vanuatu (Brouard & Grandperrin 1985) and PNG (Richards & Sundberg 1984).
Brouard & Grandperrin (1985) concluded that there was little difference between day and night catch rates, but that when the data were broken down by 40m depth
intervals, catch rates were greatest in shallow water at night. Richards & Sundberg (1984) reported on a study that was specifically designed to test the differences in
CPUE among depths and times of day. They found that peaks in CPUE occurred progressively later in the day from the deepest to the shallowest depths over 24 h.
Dalzell & Preston (1992) summarized the information from surveys of deepslope stocks to investigate the interaction of depth and time of day on deepslope catches.
CPUE increased with increasing depth between 50 and 150m, was constant between
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150 and 250m, and declined with increasing depth between 250 and 450m. Hourly catch rates showed no particular pattern in shallow waters, but resolved into peaks
at 12.00 and between 22.00 and 02.00h at depths beyond 200m. The average pattern of catch rates over a 24h period, for all depths, produced peaks at 12.00,
15.00, 20.00 and 03.00h. Ralston et al. (1986) found that the abundance of fish at Johnson Atoll peaked at 170m depth with a smaller minor peak at 250m. Overall,
the pattern of abundance was similar to the depth pattern of catch rates observed from survey fishing data by Dalzell & Preston (1992). Ralston et al. (1986) found that
on average, catch rates were twice as high in the morning than in the afternoon but that catch rate began to improve with the onset of the evening crepuscular period.
The foregoing data do not include information on individual species captured and the resultant patterns in CPUE with depth and time of day are the result of various
species interactions. Haight et al. (1993) reported the results of experimental fishing on the outer banks of the Hawaiian Islands for eteline snappers using handlines
and longlines. This study showed that overall CPUE in numbers of fish fluctuated throughout the diel cycle and peaked during the crepuscular periods (04.00 to
06.00h and 18.00 to 20.00h). Catch rates of individual species also fluctuated markedly, with Pristipomoides spp. and Etelis coruscans being caught mainly in the
morning, in contrast to E. carbunculus, which was captured mainly during the evening and night. Other species, such as Aprion virescens, were captured throughout
the day but generally not at night.

Unlike shallow water reef fishes, there has been a considerable amount of stock assessment research and biological studies made on Pacific deepslope fish stocks. A
series of review papers on deepslope fish biology and stock assessment was edited by Polovina & Ralston (1987). Age, growth and related biological studies
commenced on deep slope species in the Pacific during the 1980s, principally on the eteline species with the intent of monitoring the effects of fishing and conducting
stock assessments. Ralston (1981) presented an account of the population biology of Pristipomoides filamentosus in the Hawaiian Islands that included a new
approach to ageing longlived fishes from daily growth increments. The same method, which has also been described in detail by Ralston & Miyamoto (1981), does
not require a complete reading of the growth record in the otolith microstructure but only requires several counts of increment densities at successive distances from the
otolith core so that these can be integrated to give a complete estimate of the age of the fish.
This methodology has been used to age several deepslope species from the Northern Mariana Islands (Ralston & Williams 1988b), PNG (Richards 1987) and
Vanuatu (Brouard & Grandperrin 1985). A comparative study of the age and growth of Etelis carbunculus from Hawaii, Northern Mariana Islands, Vanuatu and
French Polynesia was made by Smith & Kostlan (1991) using Ralston’s technique. They were able to show that there were major differences in age at length for the
four widely separated stocks of the same species. Other estimates of growth of deepslope species have been made by analysis of length frequency data from Vanuatu
(Carlot 1990) and Tonga (Sua 1990, Mees 1994). Most of the studies listed above have computed mortality rates for deepslope species from length frequencies,
catch curves or a combination thereof. Deepslope species typically have life spans of between 20–30yr, with concomitantly low natural mortality rates. Ralston (1987)
has reviewed the mortality rates of deepslope snappers and groupers and has suggested that agents responsible for natural deaths in these species include predation,
parasitism, cold
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water shock and red tide poisoning. Ralston (1987) has also reviewed information from fisheries for tropical deepslope snapper and grouper species and concluded
that these species have a relatively limited productive capacity and are vulnerable to overfishing.
Two basic approaches have been used to estimate the MSY from deepslope fish stocks in the South Pacific: multispecies surplus production models where the
catch of all or a group of the commonest species are combined and plotted against effort; and biomass estimation and yield calculations based on biological
characteristics. A third method for assessing deep slope stocks is to conduct direct observations on fish populations in situ while conducting survey fishing. Such a
series of observations has been made by Ralston et al. (1986) at Johnson Atoll, where survey fishing was complemented by a series of deepslope fish abundance
estimates made from a small submersible. More recently, Ellis & DeMartini (1994) have correlated estimates of abundance of juvenile Pristipomoides filamentosus
and other Hawaiian deepslope fish made by a remote video camera with demersal longline catch rates targeted at the same stocks. However, such methodology,
requiring expensive specialized technologies and skills, is unlikely to be commonly used in the Pacific.
Ralston & Polovina (1982) fitted a multispecies version of the Schaefer stock production model to catch and fishing effort data from a deepslope handline fishery
on the banks between the islands of Maui, Lanai, Kahoolawe and Molokai (Fig. 2, 57) in Hawaii. They found that the annual predicted MSY was 106t or
272kgn.mi−1 of 100 fathom isobath. Ralston & Polovina (1982) explained that expressing yield per linear measure was probably more appropriate for steepsided
islands than the use of an areal or planar measure. Other authors have adopted this convention and expressed yields as kgn.mi−1 of 100fathom or 200m isobath.
Elsewhere, King (1992a) fitted simple Fox and Schaefer production curves to data for catches of the five principal species (P. filamentosus, Etelis carbunculus, E.
coruscans, Epinephelus septemfasciatus and E. morrhua) in the Tongan deep slope fishery. King (1992a) found that the MSY predicted by the two models was
255 and 284tyr−1 respectively. A similar analysis for all demersal species suggested MSYs of 400 and 560t respectively, but the fit to the data was rather poor. The
length of the 200m isobath in Tonga is estimated to be 294n.mi (Latu & Tulua 1991) thus the yields estimated from King’s (1992) analysis range from 0.87 to
0.97tn.mi−1 for the principal five species to between 1.36 and 1.90tn.mi−1 for the total demersal catch.
The other approach to estimating MSY from deepslope stocks in the South Pacific is to use depletion models to estimate the unexploited biomass and then, using
the biological characteristics of the stock, estimate what fraction of the virgin biomass can be harvested. The simplest approach has been to conduct shortterm
intensive fishing experiments to generate cumulative catch and CPUE data for use with a Leslie stock depletion model (see p. 420). For situations where fishing has
commenced and longer time series of catch and effort data are available, then Allen’s (1966) method, which incorporates natural mortality rates and recruitment, is
more appropriate.
Polovina (1986) used the simple shortterm Leslie depletion method to estimate the biomass and catchability coefficients of deepslope fishes in the 175 to 275m
depth range, from a 13day intensive fishing experiment at a small pinnacle reef in the Northern Mariana Islands. Based on these results and with further fishing at most
of the islands and seamounts in the Northern Mariana Islands, Polovina & Ralston (1986) were able to estimate the total biomass of deepslope fishes in this
archipelago and the MSYs for the seven principal species in the catch, namely, one jack, Caranx lugubris, and six snappers, Pristipomoides zonatus, P. auricilla,
P. filamentosus, P. flavipinnis, Etelis carbunculus, and E. coruscans. They suggested that the MSY for the deepslope stocks of Northern Mariana Islands was
about
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onethird of the original unexploited biomass, which ranged between 0.260 and 1.207tn.mi−1 of 200m isobath with a mean of 0.675tn.mi−1 200m isobath, giving an
MSY of 0.22tn.mi−1 of 200m isobath or an absolute value of 109t.
The more complex approach to estimating biomass and yield using Allen’s method has been employed by Langi et al. (1988) for deep slope fisheries on seamounts
around the Tongan archipelago. They stated that the average surplus production or MSY from three seamounts was 0.74tn.mi−1 of 200m isobath, or an absolute value
of 217t for the 294n.mi of 200m isobath in Tongan waters. The Allen and Leslie depletion methods were both used to estimate biomass and MSY for several other
locations in the South Pacific and are presented in a series of papers contained in Polovina & Shomura (1990). These locations include banks and seamounts in Fiji,
island slopes in Vanuatu and island slopes and seamounts in PNG, as well as a reanalysis of the data from Tonga using an extended data set. A summary of these
estimates was presented in Polovina & Shomura (1990), which suggested that the unexploited recruited biomass ranged from 0.2 to 7.0tn.mi−1 of 200m isobath, and
that MSY lay in the range of onetenth to onethird of the unexploited virgin biomass.
Mees (1994) reanalyzed catch and length frequency data from the Tonga deepslope fishery and fitted a dynamic production model to catch and effort data from
the Tongan fishery to obtain a total MSY of 588t. Using a modification of Allen’s model, Mees (1994) estimated a yield of 0.50–0.77tn.mi−1 of 200m isobath for a
guild of six main species in the fishery (Pristipomoides filamentosus, P. flavipinnis, Etelis carbunculus, E. coruscans, Epinephelus septemfasciatus and E.
morrhua), and of 0.33–0.63tn.mi−1 of 200m isobath for Pristipomoides filamentosus only. Mees (1994) noted that individual species’ catch rates showed an
increase over time for Etelis coruscans and a decrease for Pristipomoides filamentosus. This was due less to depletion, however, than an increasing trend to fish
deeper to target for E. coruscans which was more valuable on export markets. Mees (1994) did conclude, however, that there might be some fishinginduced effects
between E. coruscans, and Epinephelus septemfasciatus, with catch rates of the former species increasing as populations of the latter are reduced. Mees (1994)
reasoned that E. septemfasciatus is the largest fish exploited in the fishery and large specimens will not be replaced rapidly. As they are fished out, the remaining
smaller specimens provide less competition to Etelis coruscans for the baited hooks and hence the catch rate of this species increases. Indeed, Mees notes that
fishermen in Tonga report actively fishing for this species in order to remove it in order to increase catches of E. coruscans.
A similar observation was made by Polovina (1986) during shortterm intensive fishing experiments in the Northern Mariana Islands. Polovina noted that as the catch
rates of Pristipomoides zonatus and Etelis carbunculus declined during the 13 day fishing experiment, the catch rate of Pristipomoides auricilla showed a marked
increase. Polovina suggested that the interaction of P. auricilla and Etelis carbunculus was unlikely to be attributable to the latter species living at greater depths.
However, species interaction would most likely occur between P. zonatus and P. auricilla that were more abundant in the same depth interval (100–120m). Polovina
(1986) reasoned that if P. zonatus was more aggressive in pursuing fish baits than P. auricilla, or in some other way affected the behaviour of the latter, then the initial
catchability of P. auricilla would be low but would rise as the population of P. zonatus was reduced. Polovina (1986) modified the simple Leslie depletion model to
account for this species interaction.
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Only one deepslope fishery of any significance (Tonga) persists in the South Pacific. Despite the initial optimism that was generated by the exploratory surveys on
virgin stocks in the Pacific, it was not immediately appreciated that these populations comprised large, slow growing species, and that most countries of the region, by
virtue of their size, had limited stocks that could withstand only moderate exploitation. In human terms this meant that deepslope fisheries must remain small and indeed
the Tongan fishery has survived only by reducing fleet size by half and maximizing the value of the catch by exporting P. filamentosus and Etelis coruscans to Hawaii.
The access to overseas markets and the importance this has played in the survival and collapse of deepslope fisheries cannot be emphasized too strongly. The same
is also true of the growing interest in catching large valuable tunas for the Japanese market (see p. 451). PNG has by far the largest resource of deepslope fish in the
region by virtue of the extent of the 200m isobath. Furthermore, these stocks have been shown to be productive and could probably generate between 500 to
1500tyr−1 at MSY (Dalzell & Preston 1992). However, PNG does not have direct air links to either Hawaii or Japan.
Domestic fish marketing in PNG is poorly developed and only people in the capital, Port Moresby, are prepared to pay relatively high prices for fresh fish.
Elsewhere fresh fish is sold for US$1.0–2.0kg−1, which does not encourage fishermen to invest in the equipment necessary for deepslope fishing. There have been a
number of schemes to transport fish to the capital and encourage fishing on reef, pelagic and deepslope stocks but these could not be operated at a profit and have
never been effective. Another factor that militates against development of deep slope and other fisheries in PNG, and indeed many other countries of the South Pacific,
is the availability of cheap fish imports such as barracoutta and hoki. These species are caught in large volumes by trawlers in New Zealand, and can be sold in bulk for
a fraction of the cost of fresh fish within the Pacific islands.
Direct air links to Hawaii or Japan are not, however, a guarantee that a deepslope fishery will be successful. Itano (1991) documented in detail the history of deep
slope fishing in American Samoa from the mid 1960s to the mid 1980s. The limited habitat area meant that catch rates in the nearshore area declined rapidly and
fishermen were obliged to venture further offshore to seamounts and banks to maintain profitable catch rates. During the early and mid 1980s fish were shipped to
markets in Hawaii via the regular air service between Honolulu and Pago Pago (Fig. 2, 77). However, delays in payments to fishermen in American Samoa acted as a
disincentive to fishermen who were already operating on a narrow profit margin. Itano (1991) states that with such a limited resource, deepslope fisheries in American
Samoa are destined to be pulse fisheries operating over short periods on a boom and bust cycle.
The ease with which deepslope stocks can be depleted is illustrated by examples of fishing on pinnacle reefs in Guam, Fiji and American Samoa. In Guam, one
fisherman fished a pinnacle reef for 17 months and drove the catch rates down to zero (Ikehara et al. 1970). Similarly, catch rates on some of the banks and seamounts
in the Fijian fishery were driven to zero or near zero levels in less than a year. In American Samoa experimental fishing between February and May 1986, on 2% Bank,
a pinnacle reef between Tutuila and Manua (Fig. 2, 53) Islands, resulted in the removal of 78% of the eteline snappers on this seamount (Moffitt 1989). Clearly,
promotion of open access commercial fisheries on these limited resources does not make sense. However, development planners promoted these strategies in countries
such as Tonga, American Samoa, Vanuatu, Fiji and Tonga by providing easy credit for the acquisition of boats and fishing gear. As a consequence, the deepslope
fisher
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ies in these countries soon became overcapitalized and most fishing operators who had taken loans to enter the fishery defaulted on repayments (Itano 1991,
T.J.H.Adams unpub. data, S.Tulua, Ministry of Fisheries, Tonga, pers. comm.). The fishery in Fiji, which was developing into a more sophisticated commercial
operation, simply shifted to targeting large tuna that could realize much better prices on overseas markets that had been established previously for deepslope fishes
(Adams 1990). The Vanuatu fishery has a sizable demand from the hotel and restaurant trade for fresh deepslope fish but many fishermen have had to target shallow
reef species to keep their fishing operations profitable.
Fishing for deepslope stocks may indeed commence again in Fiji, and be encouraged in PNG, if market demand for deepslope species improves and markets
open in other neighbouring countries such as New Zealand, Australia and Southeast Asia. There are indications that deepslope fishing is expanding in the Solomon
Islands and may offer excellent prospects for domestic and international markets (P.Cusack, South Pacific Commission, pers. comm.). However, it is unlikely that there
will be another major expansion of fishing for deepslope stocks comparable with the increase in effort during the 1980s, particularly where there is now the possibility
of fishing profitably for large pelagic fishes (see p. 451).
Nearshore pelagic fisheries
Large pelagic species
Description
Large pelagic fishes, including coastal and offshore species, have always been important in the subsistence diet of Pacific islanders. Tuna forms the basis of artisanal
fisheries in Kiribati, PNG and French Polynesia, and other large pelagic species, such as Spanish mackerel (Scomberomorus commerson), are important components
of commercial fisheries in the Western Pacific. Perhaps the commonest method of catching large pelagic fishes is trolling with lures or baits. Other methods of fishing for
large pelagic fishes include midwater handlining and poleandline fishing for tuna.
Originally, troll fishermen would fish along the edge of the reef or search for schools of pelagic fishes feeding on the surface and troll around these. More recently,
troll fishermen and handline fishermen in the Pacific have been increasingly fishing around anchored rafts or buoys, deployed with the objective of aggregating pelagic
fishes. Fishermen in all parts of the tropics have known that objects floating on the water surface such as logs, trees, large seaweed mats and even whales and whale
sharks may have associated schools of tunas and other pelagic species. For many years, therefore fishermen in Southeast Asian countries such as the Philippines have
used this behaviour of pelagic fishes by deploying rafts or fish aggregating devices (FADS) to concentrate schools for capture.
This technology began to be used increasingly with fleets of tuna purseseiners and poleandline fishing vessels in the Western Pacific. It was realized that FADS
could benefit smallscale artisanal fishermen in the coastal zone. These rafts when anchored in relatively shallow water will aggregate coastal species such as Spanish
mackerels, mackerel and bullet tuna, trevallies and rainbow runners. FADS deployed in deeper waters will concentrate schools of oceanic tuna such as skipjack,
yellowfin and bigeye, and billfishes such as marlins, sailfish and swordfish. FADS can be constructed from a variety of materials. In Southeast Asia, FADS are often
made from floating bamboo rafts with coconut palm fronds suspended beneath.
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More modern designs of FADS have been deployed at various locations in the South Pacific and various raft designs and mooring systems are given in Boy & Smith
(1984).
In some Pacific island countries the dropoff from the shelf to the abyssal zone can be very close to shore and fishermen may have a tradition of catching oceanic
pelagics, either by trolling or handline fishing. Cusack (1987) describes a method of midwater handline fishing common in the islands of Polynesia and Micronesia,
where a hook baited with a small fish, such as a scad, is wrapped around a flat stone and secured with a slipknot. Pieces of bait are fixed to the stone with a leaf or
piece of bait skin. The stone and bait assembly sink rapidly and the fishing line is allowed to run freely without tension. When the required depth is reached the
fisherman applies tension to the line to release the slipknot, allowing the chum to disperse and presenting the unencumbered baited hook. The dropstone rig is fished
at about 25m to catch rainbow runners (Elagatis bipinnulatus) and as deep as 150–180m to catch yellowfin and bigeye tuna.
Poleandline fishing in the South Pacific is usually associated with catching skipjack on the open ocean in large (>60gr.t) fishing boats. However, a smallscale pole
andline fishery for skipjack tuna operates within the near shore waters of Tahiti (Society Islands) to supply the domestic fish market (Chapman & Cusack 1988a).
Traditionally, poleandline fishing was an important communal fishery where large double canoes were equipped with baskets lashed alongside to keep live bait. The
bait fishes were broadcast on the water to attract feeding schools of skipjack within range of the motherofpearl lures. With social and economic change and the
introduction of modern fishing materials and motorized vessels, the fishery has become a highly developed and competitive local industry to supply fresh tuna to the
local markets. Live bait is no longer used in the modern fishery, that uses halfcabin launches about 10m in length, powered by large diesel engines ranging from 200 to
375hp. The gear used in this fishery consists of a long bamboo or fibreglass pole, about 7m in length, with a length of nylon monofilament line attached to which is a
pearlshell lure with a barbless hook. The tuna boats, or bonitiers, are driven at high speed to chase skipjack schools feeding on bait fishes. The objective is to get
ahead of the school and anticipate the direction it will move so that it approaches the bonitier from the stern in order that poling can commence. The pearlshell lures are
trailed in the water as the boat moves slowly ahead of the school. When a strike is made the fishermen uses his body weight to heave the fish from the water into the
boat. Once an adequate volume of tuna has been caught the bonitiers run at high speed for Tahiti to sell their catch in the afternoon market in Papeete.
A smaller nearshore canoebased poleandline fishery for mackerel tuna (Euthynnus affinis) at Timoenai Island (Fig. 2, 102), PNG, has been described by Haines
& Chapau (1991). Live bait are caught around reefs and mangrove creeks by drivein net fishing and kept alive in a basket or the flooded hull of a canoe. The main
species of bait fishes are cardinalfishes (Apogonidae). Fishing poles are made from sago palm fronds used with barbless wire hooks. Haines & Chapau (1991)
reported that each fishing trip lasts about 4–5h, with fishermen preferring to fish at dawn or dusk. The slow speed of the canoe means that the fishermen can only
approach schools that surface close to the fishing vessel and cannot chase schools as they move away.
Troll fishing from small skiffs and dinghies is a common fishing method throughout the Pacific and the various techniques employed are summarized in Preston et al.
(1987). In some locations, small pelagic fishes such as halfbeaks or flying fishes are favoured bait and there are specialized fisheries to catch these for bait and food. In
the Belep (Fig. 2, 8) islands of New Caledonia, fishermen both troll and handline for Spanish mackerels. Handlines are baited with sardines caught usually by cast
netting on the beach. When not trolling, the fish
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ermen let out surface handlines baited with sardines, and drift with currents in the lagoon waiting for a strike. Fishing is conducted from small skiffs 4–5m in length and
this fishery catches annually about 15t of Spanish mackerel which are smoked, canned or sold fresh in the capital Noumea (Chapman & Cusack 1988b).
Spanish mackerel also dominates troll catches in the smallscale, reef associated fishery in the Tigak Islands, northern PNG (Wright & Richards 1985). As in most
other Pacific islands, trolling in the Tigak Islands is conducted from small dinghies and powered canoes with island fishermen using halfbeaks (Hemiramphidae) as
trolling bait. Troll fishing in the Tigak Islands is usually not a fulltime activity, but is part of a repertoire of fishing methods that may be deployed during a fishing trip that
also includes bottom handlining and spear fishing. Troll fishing increases in frequency in the Tigak Islands during the middle of the year as Spanish mackerel aggregate
to spawn and hence the catchability increases.
Spanish mackerel is also the principal target for troll fishermen in Fiji and landings of this species probably exceed 375t annually (Lewis 1988a, Anon. 1994b).
Trolling is conducted along the reef edge employing a wide variety of lures and live bait. Livebait fishing for Spanish mackerel is carried out seasonally in the northwest
of Viti Levu and western Vanua Levu (Fig. 2, 110). Sardines and mackerels are caught at night in gill nets and used as live bait for drift handline fishing. Troll fishing is
the most popular method of small boat fishing around Guam and was traditionally practised from manpowered canoes prior to the advent of powered dinghies and
skiffs. The major species taken around Guam are mahimahi (Coryphaena hippurus), wahoo, skipjack and yellowfin tunas (Fig. 2, Amesbury et al. 1986). The total
troll catch around Guam varies but is usually greatest between FebruaryApril. As with locations such as New Caledonia, a large proportion of the total large pelagic
fish catch comes from recreational fishermen.
Artisanal fishermen operating from Daugo Island near Port Moresby, PNG have traditionally trolled for pelagic fishes along the outer edge of the Papuan Barrier
Reef, targeting tuna, Spanish mackerel, jacks and longtoms (Belonidae). Lock (1986a) reported that fishermen adjusted their trolling techniques to take advantage of
the seasonal abundance of tuna and longtoms. Most fishermen would also include other fishing techniques to catch reef fish such as netting, handlining and spear
fishing. The deployment of a FAD on the outer reef slope (Beverly & Cusack 1993), and the resulting abundance of large pelagic fishes, has led to more fishermen
concentrating on troll fishing (Dalzell 1993a). Fishing is conducted from fibreglass dinghies with four to six troll lines strung from a boom at the front of the vessel using
homemade and commercial trolling lures. The productivity of troll fishing around the FAD has led to a decrease in the number of fishermen targeting coral reef fishes,
and presumably to a decrease in the fishing mortality of reef fishes.
The deployment of the FAD in Port Moresby has also been of additional benefit to the game fishing club, which targets large pelagic fishes, more for sport and
trophies than for food. In some areas of the Pacific the recreational component of fisheries production can be quite substantial. In New Caledonia, for example, the
recreational catch from the nearshore area is thought to be in the vicinity of 2000t, or about half of the total coastal fisheries production (Anon. 1994a). A substantial
portion of the recreational catch is likely to comprise large pelagic species such as tuna, Spanish mackerel, wahoo and billfishes. In Vanuatu, the local game fishing club
in Port Vila is one of the major suppliers of large pelagic fishes for the domestic fish market (Anon. 1992b) as local fishermen are not experienced in the capture of
coastal pelagic species through trolling.
Large pelagic fishes form the principal targets of fishermen from small high islands lacking any substantial reef and lagoon areas, with catches taken mainly by trolling
(Dalzell &
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Debao 1994). Twothirds of the total landings on Nauru are from trolling, with the catch made up mainly from skipjack, yellowfin tuna and wahoo (Acanthocybium
solandri). Trolling is conducted from small skiffs deploying on average two troll lines and fishing mainly in the early morning and late afternoon. Although most of the
catch might be classed as recreational and subsistence fishing, between half and twothirds is sold locally. About half of the total fisheries production on Niue was
estimated to come from fishing large pelagic species by trolling and handlining (Dalzell et al. 1993). Troll fishing is conducted from small skiffs fishing in open water or
around FADS. Some Niuean fishermen also fish with midwater handlines, or the paluahi method, from canoes in open water in areas where large yellowfin and bigeye
tunas are known to aggregate, usually around headlands. This method was still very popular until recently in Rarotonga, Cook Islands, for catching large tuna relatively
close to shore. FADS have been regularly deployed around Rarotonga since 1980 and have promoted greater catches from troll fishing (Sims 1992a).
Catch composition
The catch composition by family for troll fishing in the Pacific islands is shown in Table 14. The tuna and related species such as wahoo and Spanish mackerel dominate
the catches in most locations. The notable exception is Fiji, where substantial amounts of barracudas and jacks are caught along with the tunalike species. In Fiji,
trolling close to reefs attracts reef predator species into the open water, hence 12% of the Fiji catch were groupers, mainly coral trout (Plectropomus spp.). Other reef
species such as snappers and emperors are also occasionally taken by trolling over the shallow reefs or along the reef edge. Overall, scombrids form about 80% of the
troll catch with between 7 and 8% each formed from jacks and barracudas. Very large pelagics such as certain sharks and billfish (Istiophoridae) are occasionally
taken, but these are more deliberately targeted by game fishermen rather than artisanal fishermen.
The scombrid component of the large pelagic catch in the major Melanesian islands of the western Pacific is usually far more varied than around the smaller islands of
the north and central Pacific. Spanish mackerel dominate troll catches in PNG, Solomon Islands, Fiji and New Caledonia. All these countries have substantial areas of
shallow lagoons fringed with extensive mangrove forests, which is nursery habitat for juvenile Spanish mackerel. In Vanuatu, the only Melanesian country that lacks
extensive mangrovefringed shallow lagoon areas, Spanish mackerel are not abundant and are only caught in a few locations. The scombrid catches in the large
Melanesian islands contain other coastal scombrids such as the island bonito (Euthynnus affinis), bullet tuna (Auxis spp.) and shark mackerel (Grammatorcynus
bicarinatus). Around the coasts of the smaller Pacific islands in the northern and central Pacific, troll catches are dominated more by skipjack (Katsuwonus pelamis)
and yellowfin tuna (Thunnus albacares). Spanish mackerel are found in Palau, the only islands with sufficient mangrove and estuarine habitats.
Catch rates
Catch rates from troll fishing for large pelagic fishes in various parts of the Pacific are shown in Table 15. Average CPUEs ranged between 1.8 and 8.8kgline−1h−1 with
an overall average of 4.5 kg line−1h−1. Table 15 also includes the dominant species in the catches and demonstrates clearly the difference between the large Melanesian
islands, where catches are dominated by large coastal pelagic species and reef associated fishes, and the smaller islands
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Table 14 Percent composition of catches made by trolling along the coastal margins in the South Pacific region.
Family Fiji PNG New Caledonia Am. Samoa Tokelau Tuvalu Nauru Palau Niue Wallis & Futuna Kiribati (Gilberts) Tonga Vanuatu Cook Islands
Scombridae 29.7 84.6 80.6 86.0 100 79.4 88.5 45.9 85.9 52.9 77.8 98.4 92.9 97.6
Sphyraenidae 33.7 10.8 2.0 1.5 1.8 38.2 1.4 18.6 1.2
Carangidae 22.4 5.8 5.2 12.7 8.2 2.9 27.1 19.4 1.7
Istiophoridae 0.2 6.0 1 0.6
Serranidae 12 1.1 0.3
Lutjanidae 2.1 1.1 3.8 2.1 1.4 2.8
Belonidae 0.2 0.8 0.6 0.3
Coryphaenidae 0.2 0.5 0.2 10.6 1.5 0.7 4 1.6
Others 0.1 9.6 0.5 6.0 11.2 0.9 0.2 0.8
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Table 15 Summary of catch rate (CPUE, catch per unit effort) and catch composition data for trolling on coastal pelagic stocks in the South Pacific
region
CPUE (kg line−1h−1)

Location Range Mean Principal species in catch Source
Papua New Guinea 3.7–6.9 4.9 Euthynnus affinis, Dalzell & Wright 1986
Auxis thazard, Wright & Richards 1985,
Scomberomorus commerson Lock 1986a, Anon. 1984b
Palau 3.7–12.6 8.2 Katsuwonus pelamis, Anon. 1991b, 1992e
Thunnus albacares,
Acanthocybium solandri
Fiji 1.6–16.2 5.7 Sphyraena queni, Chapman & Lewis 1982,
S. barracuda, Lewis et al. 1983
S. commerson,
Plectropomus sp.
Tuvalu 0.5–7.0 2.7 A. solandri, Chapman & Cusack 1988c
K. pelamis,
T. albacares,
Istiophorus platypterus
New Caledonia 0.0–7.3 4.0 S. commerson, Chapman & Cusack 1988b
E. affinis,
A. solandri,
T. albacares
Tokelau 0.0–68.9 8.8 K. pelamis, Taumaia & Cusack 1990
T. albacares
Nauru 0.0–20.0 4.7 K. pelamis, Dalzell & Debao 1994
T, albacares,
Elagatis bipinnulatus,
Sphyraenidae
Niue 1.3–3.8 2.4 A. solandri, Dalzell et al. 1993
T. albacares,
K. pelamis
Am. Samoa 0.5–6.1 3.2 K. pelamis, Buckley et al. 1989
T. albacares, Craig et al. 1993
Makaira mazara,
Coryphaena hippurus
Wallis & Futuna na 6.0 K. pelamis, Taumaia & Cusack 1988
T. albacares,
Sphyraena queni
Kiribati 0.5–8.0 3.8 T. albacares, Taumaia & Cusack 1989
K. pelamis,
E. bipinnulatus
Vanuatu 0.51–14.0 1.8 T. albacares, Cillauren 1988
K. pelamis,
C. hippurus
Cook Islands 0–12.3 1.8 K. pelamis, Chapman & Cusack 1988d
T. albacares
of the Pacific where oceanic pelagics dominate the catch.

Catches of large pelagic fishes are strongly seasonal, with seasons becoming more pronounced at higher latitudes. Detailed information on catch seasonality is
available from Guam in the northern hemisphere of the Pacific (13°N) (Amesbury & Babin 1990) and Niue in the south (19°S) (Dalzell et al. 1993). More species
were included in the study in Guam by Amesbury & Babin (1990) but catch data for skipjack, yellowfin and wahoo were included in both studies. Temperature studies
at Guam show that there is a yearround permanent thermocline between 120 and 400m in which temperature drops from 28°C to 7°C. In winter, when surface waters
are coolest, there is a deep mixed layer from the surface to 120m. During spring and summer the depth of the mixed layer is reduced and a seasonal thermocline
develops before reestablishment of the deep mixed layer with the onset of autumn.
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According to Amesbury & Babin (1990) catch rates for skipjack and yellowfin were greater during the summer months when surface waters were warm and the
seasonal thermocline had developed. Catch rates for wahoo demonstrated a clear peak in the winter months when the surface waters were cooler and the mixed layer
deepest. Catch rates of wahoo south of the equator were highest during the austral winter, between June and August. Yellowfin catches were also greatest in the winter
months, in contrast to the seasonal pattern in Guam, while skipjack catch rates peaked in the summer months in common with the seasonally for the same species north
of the equator. Other species included in the Guam study were blue marlin (Makaira mazara) and dolphin or mahimahi (Coryphaena hippurus). The CPUE for
marlin was similar to that of the two tuna species with peak catch rates in summer, while the dolphin CPUE showed a marked peak during the cooler winter months.

The biology and stock assessment of skipjack, yellowfin, bigeye (Thunnus obesus) and albacore (T. alalunga) have been studied both nationally and regionally in the
Pacific. The South Pacific Commission has mounted major regional studies of all four species over the past 20yr which has resulted in a series of publications on the
biology and stock assessment of these species. Much of the information on the population biology of these species has come from largescale tagging projects
conducted throughout the South Pacific by the South Pacific Commission in cooperation with national fisheries departments and other scientific agencies. The most
recent reviews of the biology and stock assessment of these four important commercial pelagic species in the South Pacific region are contained in Shomura et al.
(1993) and the current status of western tropical Pacific tuna fisheries is summarized in Anon. (1995).
In this paper we are concerned more with the large pelagic species that are confined to the coastal margins. These species have received much less attention in the
South Pacific, although they have been the objects of considerable study in South and Southeast Asia where they form the basis of large industrial fisheries. Spanish
mackerel populations along the GBR have been studied in detail and information on age, growth, reproduction, early life history and feeding habits are contained in
Munro (1942) and McPherson (1987, 1992, 1993). Less information on this species is available from the Pacific islands. Shaklee et al. (1990) examined the stock
structure of Spanish mackerel from eastern and western Australia, Fiji and PNG using electrophoretic techniques. Fish to the southeast of Torres Strait form an
Australian east coast stock, whereas fish in Torres Strait and the Gulf of Carpentaria (Fig. 2, 120) are part of a northern Australian stock that extends from PNG to the
western coast of the continent. Populations from northern PNG (New Ireland) and Fiji were found to be distinct stocks that were effectively isolated.
Lewis (1975) presented some notes on the biology and fisheries for Spanish mackerel in PNG waters and summarized the available information on behaviour,
reproduction, spawning sites and feeding habits. Similar information on Spanish mackerel in Fiji is included in a study of several large coastal pelagic species by Lewis
et al. (1983), who reported that Spanish mackerel in Fiji appeared to grow more slowly than specimens sampled from the GBR. Lewis et al. (1983) provided
biological information on a range of other large coastal pelagic species including shark mackerel (Grammatorcynus bicarinatus), dogtooth tuna (Gymnosarda
unicolor), barracudas (Sphyraena barracuda and S. queni) and trevallies (Caranx ignobilis, C. melampygus and C. papuensis). Information was given on size
distribution, spawning season, size at first maturity and ecology of these species.
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The biology of the trevallies C. melampygus and C. ignobilis from Hawaii were studied in some detail by Sudekum (1984). Sudekum was able to estimate the age
and growth of C. melampygus from otolith microstructure but gave no details of ageing the other species in the study. Other information presented by Sudekum for
these two species includes reproductive and feeding biology. A synopsis of biological data on the amberjack (Seriola dumerilii) in Hawaiian waters is given in Uchida
& Uchiyama (1986), including information on age and growth. This species ranges from surface waters to depths of up to 250m and can frequently be caught by deep
slope handlining as well as by trolling.
Other large coastal pelagic species for which some biological information is given in Uchida & Uchiyama (1986) include dolphin (Coryphaena hippurus), wahoo
and the mackerel tuna or kawakawa (Euthynnus affinis). The latter species is very common throughout most of the coastal waters of the Pacific, as are the small
bullet tuna (Auxis thazard and A. rochei). Little of the biology of these three inshore scombrids is documented for stocks in the Pacific islands. Wilson (1981a)
presents a few notes on the biology of Euthynnus affinis from PNG and northern Australia and Lewis et al. (1983) provide similar information for this species in Fiji.
A few observations on the biology of E. affinis are also given by Haines & Chapau (1991) from specimens captured in Manus, northern PNG, while Mobiha (1993)
has recorded the spawning seasonally of both E. affinis and Auxis thazard from the waters around New Ireland, also in northern PNG. Wilson (1981b) gives an
account of the biology of the longtail tuna (Thunnus tonggol), a species more commonly associated with extensive shelf zones and found in the Gulf of Papua, where it
is taken by gill netters targeting sharks. Wilson provides information on age and growth, from tagging and otolith microstructure, reproductive biology and stock
structure around PNG and Australia.
One of the most notable developments of the last 5 years in the tropical western Pacific has been the development of domestic longline fisheries for fresh, chilled
yellowfin and bigeye tuna to satisfy the Japanese demand for raw fish or sashimi. It was the introduction of monofilament longlines in the Hawaiian longline fishery in
the early 1980s that led to improved catches of these species (Cook 1989). Deployment of monofilament line can be automated with a line thrower, allowing steeper
catenary longline curves that let this gear fish deeper than kuralon longlines, and this longlining method spread rapidly to other parts of the Pacific. Improved airline
connections between the Pacific islands and Japan, Hawaii and the continental United States has meant that domestic commercial longline fisheries have developed (or
are developing) in American Samoa, Fiji, Palau, Federated States of Micronesia, Marshall Islands, Guam, French Polynesia, New Caledonia and Tonga. A major
limiting factor for such fisheries is access to markets, which is dictated mainly by airline schedules and the space available for cargo on commercial aircraft. Catches by
these smallscale domestic longline tuna fisheries are classified by the South Pacific Commission’s Tuna and Billfish Assessment Programme in the Regional Tuna
Bulletins, and are not included in the catch summaries in Appendix 1 (p. 510) and Tables 26 and 27 (p. 499). However, this fishery overlaps significantly with coastal
fisheries, particularly where vessels are used for a variety of fishing methods.
In parallel with the development of the smallscale, longline fishery has been experimental fishing for small pelagic stocks to supply bait for longlining operations.
Longlining in the tropical Pacific has relied for the most part on imported bait from Japan. Partial or total replacement of imported bait could decrease operating costs
and possibly increase the spread
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of economic benefits from longline fishing. Small pelagic stocks (scads, sardines and herrings) have traditionally featured in the subsistence diet of many Pacific islands
and are caught for sale in domestic markets (see also p. 463). Experimental bait fisheries using purseseines and lift nets have been established in Tonga (King 1993a)
and the Federated States of Micronesia (Anon. 1992c), while the potential for local bait supply was investigated in PNG (Dalzell 1993a). Locally available small
pelagic species have been briefly tested as longline bait by commercial boats in Fiji, but were too seasonal and reliable supplies were difficult to obtain (Adams unpub.
data). Longline fishermen are notoriously conservative in their choice of bait. Experimentation is expensive, and local substitutes are likely to be accepted only if good
catch rates can be consistently demonstrated.
A further development has been the adoption of a technique first pioneered in Indonesia where live milkfish are used to bait longlines thereby markedly improving
catches. Plans have been made to develop milkfish culture in Chuuk by a longline fishing company based in the Federated States of Micronesia (Lindsay 1994).
Milkfish culture is a tradition in Micronesia and may offer further potential for fishermen to share in the economic benefits of longline fishing.
Greater emphasis is now given to developing nearshore pelagic fisheries to improve fishermen’s incomes, relieve fishing pressure on reef and deepslope stocks and
to improve the supply of fresh fish on domestic markets. In a few areas, such as Port Moresby, Rarotonga (Cook Islands), Apia (Western Samoa) and parts of Vanua
Levu (Fiji) there are now some commercial fishing communities that are entirely dependent on FADS for their livelihood. Artisanal fishermen may also receive further
economic benefit if they are able to catch highvalue sashimigrade tuna and have a local buyer with access to overseas markets, as in Suva. In urban areas where there
is a substantial demand for fish, such as Port Moresby and Apia, the deployment of coastal FADS has been shown to be very successful, and fishermen are able to
make substantial earnings from large pelagic catches (Watt & Cusack 1992, Beverly & Cusack 1993).
Small pelagic species
Description
Small pelagic fishes in the South Pacific are important for subsistence and smallscale commercial fisheries. In some areas, where coral reefs are not abundant, small
pelagic fish may be a staple source of protein. Small pelagic fishes as defined here refer to mackerellike and herringlike fishes that rarely exceed 500 g in weight and
more typically range in weight from 5 to 100 g. They include small clupeoid fishes such as the anchovies (Engraulidae), sprats, herrings and sardines (Clupeidae), small
mackerels of the genus Rastrelliger, round scads and bigeye scads (Carangidae), halfbeaks and flying fishes (Hemiramphidae and Exocoetidae), hardyheads
(Atherinidae) and fusiliers (Caesionidae).
The taxonomy and identification of small pelagic fishes is problematic, given the close similarities among species and the existence of congeneric pairs of cooccurring
species with very similar ecology, e.g. Encrasicholina devisi and E. heteroloba; Spratelloides gracilis and S. lewisi; Amblygaster sirm and A. clupeoides; Selar
crumenophthalmus and S. boops. Furthermore, as Lewis (1990) pointed out, other genera, such as Herklotsichthys and Dussumieria, contain separate morphs that
are likely to prove to be sibling species and is suggestive of finescale habitat partitioning. The taxonomy of the clupeoids has been stabilized to some extent by the
publication of the FAO species catalogue on these fishes
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(Whitehead 1985, Whitehead et al. 1988). Similarly, the FAO species identification sheets for the Western Indian Ocean (Fischer & Bianchi 1984), provide
reasonable accounts of the nonclupeoid small pelagic species. Some groups of fishes, such as the flying fishes, will continue to provide problems for researchers in the
region and will require further studies before these difficulties are resolved.
Artisanal fishing methods for small pelagic species include gill nets, drivein nets, fish corrals, cast nets, scoop nets and handlines. Encircling seine nets, traditionally
made from bush materials, are used in the Admiralty Islands of PNG to capture halfbeaks (Hemiramphidae) and other larger pelagics such as trevallies (Carangidae)
and mullet (Mugilidae). Gill nets are used in Fiji both actively and passively to catch schools of the mackerels Rastrelliger kanagurta and R. brachysoma. One of the
commonest small pelagic fishes found throughout the South Pacific is the bigeye scad (Selar crumenophthalmus and S. boops). In PNG, bigeye scads are caught by
gillnetting, beach seining, scoop netting and handline fishing (Hulo 1984, Dalzell 1993b). Gill nets are also used to catch Selar spp. in Palau (Johannes 1981) and in
Guam bigeye scads are captured inshore both by a variety of gill nets and by hookandline fishing (Amesbury et al. 1986). In Guam, seasonal catches of bigeye scad
occur at times when they move inshore where they are a popular target for recreational fishing. The fish captured in the inshore areas are juveniles, however, and the
larger adult fish are being caught further offshore in deeper water by using light attraction at night in conjunction with feather jigs. The bigeye scad is highly prized in
French Polynesia and is captured by drivein seine netting, gill netting, handlining with feather lures, and in fish corrals (Bagnis et al. 1974, Grand 1985, Morize 1985).
In Hawaii, a rather more sophisticated fishery has developed for bigeye scads using spotter planes to direct boats to schools for capturing the fish with surround nets
(Shiota 1986).
The goldspot herring, Herklotsichthys quadrimaculatus, is a popular target species for artisanal fishermen throughout its range in the South Pacific. In Palau this
species is caught by cast netting when it aggregates to spawn (Johannes 1981). Cast netting is also used to catch this herring in Tarawa (Kiribati) where it forms part of
the subsistence catch (Mees & Yeeting 1986). In the North Solomon Islands (Fig. 2, 70) of PNG, fishermen herd schools of goldspot herring into shallow areas
where they are caught with scoop nets and thrown into baskets carried on the back (Hulo 1984).
Scoop nets are also used in the North Solomons to capture flying fishes (Hulo 1984). The fishermen attract the flying fishes on a moonless night by a kerosene
pressure lamp or electric lamp mounted on a canoe. This type of fishing is found also in the Micronesian islands of Palau (Johannes 1981), Kiribati (Mees 1985) and
most of Polynesia where it is commonly practised. In both French Polynesia and the Cook Islands small generators are used to power electric lights. A highpowered
light is fixed to a helmet worn by the fishermen, allowing both hands to remain free to drive the boat and manipulate the scoop net (Gillett & Ianelli 1991). The boats in
these areas are designed to be driven from the bow to facilitate scooping the fish and pursuing them at high speeds over the sea surface. Flying fishes may also be
caught in gill nets and a description of gill netting for flying fishes in the Gilbert Islands of Kiribati is given by Tebano & Tabe (1993). Gill nets are set outside the reef or
in reef passages and schools of flying fishes are then driven towards the net by fishermen in either skiffs or canoes.
In Tahiti, the round scad Decapterus macarellus is caught by hookandline using coconut pulp as bait (Bagnis et al. 1974) and by netting (presumably gill netting)
in reef passes of the Leeward Island. Round scads, particularly D. macarellus, are caught by commercial fishermen in Hawaii using handlines, surround seines and
hoop nets. Gillett (1987) reports the
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successful adaptation of the Hawaiian hoop netting technique for round scads around the island of Niue. In the Cook Islands, round scads are chummed to the surface
by divers who spit out clouds of chewed coconut flesh, and are then caught with small fishing poles (P. Cusack, South Pacific Commission, pers. comm.). Chewed
coconut bait is used to chum round scads to the surface in Niue. Canoe fishermen spit the chewed coconut pulp onto the water’s surface then create an eddy with a
paddle blade to concentrate the bait. As the scads feed on the chum the canoe fishermen fish for the scads with a short bifurcated fishing pole from which two lines are
hung.
Small pelagic fishes are caught in large quantities in two industrial near shore fisheries in the South Pacific: by shrimp trawlers in Papua New Guinea and in baitfish
catches at several locations in the Pacific. The trawl fisheries in the Gulf of Papua and South Papuan coast, although targeting penaeid shrimps, catch mainly finfish.
Catches are made by 150gr.t trawlers deploying two 12fathom (footrope) nets, fishing 24hday−1 on cruises of 5–6wk duration. Dalzell (1993b) estimated that the
finfish bycatch from the Gulf of Papua fishery was about 14000t, of which about 2000t comprised anchovies (1400t) and sardines (600t). Nearly all of the fish by
catch from the trawl fishery is discarded, with only a few marketable species retained for sale in Port Moresby. Species of little commercial value, such as the anchovies
and sardines are simply thrown back into the sea.
The supportive bait fisheries for poleandline tuna fishing are conducted at night, using powerful submersible lights to attract the schools of anchovies and sprats.
Each poleandline vessel has two or three skiffs with generators from which lights are suspended, as well as from the poleandline vessel. This gives the vessel the
chance to make three to four hauls of the net to fill up the bait tanks. Hauls are usually made between 02.00 and 04.00h prior to the poleandline boat heading out on
to the open sea to catch tuna. When a haul is made the bait lamp is dimmed and drawn to the surface, which causes the bait school to concentrate around the lamp.
The lift net is then deployed to trap the bait fish. The net is lifted until the bait fish are concentrated in one corner of the net and transferred to the bait wells in the tuna
vessel by buckets. Catches are measured in numbers of buckets but can be converted to weight by a predetermined conversion factor.
As stated previously, development of longline fisheries for large pelagic fishes has generated interest in domestic production of longline bait from local stocks of
small pelagic fishes in Tonga and the Federated States of Micronesia. Trial fishing for round scads and sardines was conducted in Vava’u during 1992 using a 1200m
purse seine with a stretched mesh size of 2.0cm. As with the poleandline bait fishery, the sardine and scad schools were first aggregated around a submersible lamp.
A similar type of survey to assess the potential of longline bait supply was conducted in Chuuk Lagoon between 1991 and 1992. In this instance a dip net was
employed, rather than a purse seine.
Like most fisheries within the Pacific islands, small pelagic fisheries are smallscale artisanal activities. Only in Hawaii has fishing for small pelagic species, namely
bigeye scads, developed into an industrial scale fishing operation. The target species of small pelagic fisheries in the South Pacific tend to be the larger species such as
scads, mackerels and herrings. Rapson (1955) reports a seasonal fishery on the South Papuan coast for “whitebait”, which he describes as postlarval anchovies
(possibly Stolephorus or Encrasicholina spp.), using mosquito net or nets made from traditional material. Other than this single reference no other accounts of
traditional fishing for anchovies and sprats was found in the literature. Daytime seining for Encrasicholina purpurea is conducted in Hawaii, where it is used as live
bait for a smallscale poleandline fishing fleet.
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Catch composition
Information on the composition of small pelagic catches from artisanal fisheries is scarce and it is difficult to condense the information into simple tabular form. The gill
net fishery in Rabaul, PNG (Table 7, p. 420) targets large schools of bigeye scads, with the result that catches comprised almost entirely one species, Selar
crumenophthalmus. Similarly, hoop netting for round scads in the coastal waters of Niue yielded only a single species in the catch, Decapterus macarellus (Gillett
1987). Handline fishing for bigeye scads in Vanuatu similarly yielded a single species, Selar crumenophthalmus (Dalzell 1992b), while fishing with similar gear around
shallow water FADS at Vava’u in northern Tonga produced not only bigeye scads, which dominated the catch (73%) but round scads (23%) and other carangid
species such as rainbow runner (Elegatis bipinnulatus) (Dalzell 1992b).
The beach seine fishing data for coral and lagoon areas reported in Table 8 (p. 421) includes clupeids, atherinids and carangids in the catches. The beach seine sets
in Rabaul targeted the same stock of bigeye scads as the gill net fishery. The other beach seine catches reported in Table 8 generally consisted of small pelagic fishes.
Catches in Kiribati were predominantly goldspot herring, the hardyhead, Pranesus pinguis (Atherinidae) and the sprat Spratelloides delicatulus. Pranesus pinguis
was the dominant species also in Vava’u, Tonga, along with another atherinid, Hypoatherina ovalau and Spratelloides delicatulus, and the latter two species also
dominated beach seine catches in the Marshall Islands. S. delicatulus was one of two species to dominate beach seine catches in Tokelau, the other being the mullet,
Liza vagiensis. Beach seine catches in French Polynesia were predominantly formed from the endemic sardine species Sardinella marquesensis.
Scoop netting for flying fishes catches a variety of species although the identity of the species captured is confounded by taxonomic problems. Gillett & Ianelli (1991)
list 29 species contained in six genera that are thought to occur in the South Pacific region, but illustrate the taxonomic problems by comparing the discrepancies
between identifications of the same sets of specimens by two experienced fish taxonomists. According to Gillett & Ianelli (1991), the six genera of flying fishes thought
to occur in the Pacific islands are: Cheilopogon, Cypselurus, Exocoetus, Hirundichthys, Paraexocoetus and Pronichthys. The species identified from flying fish
catches from Tonga, Tahiti, Cook Islands, Tokelau and Tuvalu include Cheilopogon spilonopterus, C. atrisignis, C. antoncichi, C. unicolor and Cypselurus
poecilopterus. These five species were the most frequently identified from catches based on the information presented by Gillett & Ianelli (1991). Other species
occurring in fishermen’s catches included Cheilopogon suttoni, C. spilonopterus, Cypselurus oligolepis, C. angusticeps, C. pitcairnensis and Paraexocoetus
brachypterus.
Kailola & Wilson (1978) list a wide range of small pelagic species taken by the shrimp trawlers in the Gulf of Papua. These include 11 sardine, pilchard and herring
species, 12 anchovy species, bigeye scads and small mackerels. Dalzell (1986) estimated that the sardines and anchovies formed about 4 and 9% respectively of the
trawl bycatch, based on Kailola & Wilson’s data, which was collected from trawls made between 1960 and 1968. Watson (1984) made detailed observations on the
trawl bycatch in 1983 and his data suggest that sardines and anchovies formed 9 and 34% respectively of the bycatch. The dominant anchovy species were
Setipinna godavari and Thryssa setirostris, while Sardinella albela dominated the sardine catch.
The most readily comparable data from fishing for small pelagic species in the South Pacific come from the poleandline bait fisheries that have operated at one time
or another at several locations in the South Pacific. In addition, the South Pacific Commission’s Skipjack Survey and Assessment Programme conducted surveys for
tuna bait fish in the
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Table 16 Species composition in percent of bait fish catches in the South Pacific region.
Bait fish species Common names Scientific names Fiji PNG Solomon Is Palau Kiribati New Caledonia
Sprats Spratelloides spp. 23.2 18.1 11.1 40.5 3.7
Sardines Amblygaster spp.,_Sardinella spp. 15.7 5.4 1.0 27.7 15.1
Herring Herklotsichthys spp., Pelona spp. 14.7 1.2 17.7
Silversides Pranesus spp, Hypoatherina spp., Atherinomorus spp. 4.3 1.2 0.3 5.3 0.3
Mackerels Rastrelliger spp. 6.3 1.1 0.3
Cardinals Rhabdamia spp., Archamia spp. 16.5 1.6 0.2 0.8
Anchovies Encrasicholina spp., Stolephorus spp., Thryssa spp. 10.3 62.6 72.4 91.0 62.5
Fusiliers Caesio spp., Pterocaesio spp. 7.7 0.5
Scads Selar spp., Decapterus spp. 1.4
Others 9.0 0.9 13 9.0 8 18.4
South Pacific region between 1977 and 1980. These surveys suggested that five species, Spratelloides gracilis, Encrasicholina devisi, E. heteroloba,
Herklotsichthys quadrimaculatus and Atherinomorus lacunosus, were both widespread and abundant. Other common species were Hypoatherina ovalau, Selar
crumenophthalmus, Amblygaster sirm, Spratelloides gracilis, and Encrasicholina punctifer. The average catch composition of the tuna bait fisheries from six
Pacific islands is shown in Table 16 (from Dalzell & Lewis 1989). At Palau, PNG, the Solomon Islands and New Caledonia, the dominant species caught are
stolephorid anchovies4, namely E. heteroloba and E. devisi. The sprats, mainly Spratelloides delicatulus, form another important species grouping, particularly in
Kiribati where they account for 40% of bait catches. Other important components of the Kiribati catch were the sardine Amblygaster sirm, and the goldspot herring.
Bait catches from Fiji are more evenly spread among the various components and no one family or species show overall dominance. As a group, the clupeoids are
the largest fraction of the catch, accounting for nearly twothirds of the total. A feature of the Fiji bait catch that separates it from others in the region is the large
proportion of demersal cardinalfishes (Apogonidae) in the bait hauls. This feature of the Fiji bait fishery began to manifest itself during the mid 1980s when poleand
line boats began fishing with larger boukeami nets that reached to the sea floor, thus increasing the catchability of demersal species such as the cardinal fishes. The
dominant cardinal fish in the Fiji bait hauls is Rhabdamia gracilis, although in total 10 apogonids are present in the bait catch (Milton et al. 1993).
The dominant species in the nearshore purseseine fishery at Vava’u, Tonga was the sardine, Amblygaster sirm, which formed 69% of catches (King 1992b). Just
over 20% of the catch consisted of the round scads, Decapterus macrosoma (13.8%) and D. macarellus (8.0%). Goldspot herring formed almost 7% of the
balance of the catch with the rest made up of other small pelagic species. The lift net fishery in Chuuk Lagoon was dominated by round scads, which made up half the
catch. A further 23% of landings came from bigeye scads with mackerels and sardines forming about 7% each of the total catch (Dalzell 1992b).
4. Originally, all the anchovies now separated into the two genera Stolephorus and Encrasicholina were contained in the single genus Stolephorus and are still known collectively as the
“stolephorid anchovies.
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Catch rates
Dalzell (1992b) quotes average catch rates of 17.6fishline−1h−1 or 2.3kgline−1h−1 for nighttime handline jigging for bigeye scads and round scads at Vava’u, Tonga
and 18.3fishline−1h−1 or 2.4kgline−1h−1 for the same method of fishing on bigeye scads at Port Vila, Vanuatu. Similar fishing for bigeye scads in Guam produced catch
rates ranging from 0–4.7kgline−1h−1, with a mean of 1.9kgline−1h−1 (Ikehara et al. 1970). Gill nets set on bigeye scad schools in Rabaul, PNG (Table 7, p.000), caught
between 0.7 and 6.7kgset−1 with an average of 3.0kg set−1, while average catch from beach seining on the same scad stock was estimated to be 350kg set−1 (Dalzell
1993a). Beach seining for bigeye scads at Kavieng, PNG (Table 8, p. 421) yielded catch rates ranging from 1.0 to 44.6 kg set−1, with an average of 17.3 kg set−1.
Beach seining for predominantly small pelagic fishes elsewhere in the Pacific (Table 8), yielded average catch rates ranging from 13.5 to 130.6kgset−1 with an overall
average of 40.0kgset−1. Catch rates from purse seining around Vava’u for sardines and round scads ranged from 0 to 2000kg night−1, with an average of 570kgnight−1
(King 1992b). The average catch per night from dip net fishing in Chuuk Lagoon was 0 to 211kgnight−1, with a mean of 74.6 kg night−1 (Dalzell 1992b). In neither
instance was there any information to suggest how many hauls were made per night.
Flying fish catches from scoop netting at a variety of locations in the South Pacific region are summarized in Table 17. Catches by weight were not reported from all
locations. Catches in number ranged from 10 to 40fishh−1 with an average of 19.3fishh−1. Weight CPUE ranged from 1.1 to 12 kgh−1 or an average of 5.0 kghr−1 for
those locations reporting it. The correlation between weight and number CPUEs is highly significant (r2=0.99) for the five values in Table 17 and where unknown,
weight CPUE of flying fishes can be obtained empirically from the product of CPUE in numbers and 0.245. For example, based on the data of Tebano & Tabe
(1993), average catch rates for flying fishes caught by gill netting from skiffs in the Gilbert Islands ranged from 0.8–9.0fish h−1, with a mean of 4.3fish h−1 or 0.2–
2.2kgh−1 and a mean of 1.05kgh−1. Similarly, the catch rates from canoe fishing with gill nets was 1.0–5.0fishh−1 with a mean of 2.5fishh−1 or 0.25–1.22kghr−1 with a
mean of 0.54kghr−1.
Catch rates for commercial bait fishing with boukeami nets in various locations in the South Pacific are shown in Table 18. These averages are given by country and
represent averages over a number of years and between several locations within each country. The conversion factor for raising catch in buckets to kilogrammes was
about 2 in all cases, except for Kiribati where a raising factor of 3 is used. This is due to the predominance of larger sized bait fish in the catches at Kiribati, such as
sardine (Amblygaster sirm) and herring. Catch rate in kgboat−1 night−1 is also given, as the number of boat nights was more consist
Table 17 Catch rates from scoopnetting for flying fishes in the South Pacific region.
Country Catch rate, no. h−1 Catch rate, kg h−1 Source

American Samoa 14.9 3.6 Gillett & Ianelli 1991
Niue 40 12 Dalzell et al. 1993
Yap (FSM) 23 Gillett & Ianelli 1991
Solomon Is 15–20 Gillett & Ianelli 1991
Fiji 10 Gillett & Ianelli 1991
Tokelau 11 1.1 Taumaia & Cusack 1990
Tonga (Vava’u & Ha’apai) 20.4 5 Rata 1992
Tonga (Vava’u) 17.7 3.8 SPC unpub. data
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Table 18 Average catch rates from tunabait fisheries in the South Pacific.
Country Years Catch rate (kg boat−1 night−1) Catch rate (kg haul−1) Source
Papua New Guinea 1970–1981 170 71.3 Dalzell 1984a
Fiji 1976–1992 117 80.2 Rawlinson & Sharma 1993
Solomon Islands 1981–1988 290 109.3 Nichols & Rawlinson 1990
Palau 1964–1974 93.3 55.1 Muller 1976
Kiribati 1982–1990 116 72 Rawlinson et al. 1992
New Caledonia 1981–1983 284 100.8 Hallier & Kulbicki 1985
ently reported as the unit of effort in some instances, with effort in hauls only available for a few years. Catch rates were 71.3–109.3kghaul−1, with an overall average
of 81.5kghaul−1. The number of hauls made per night depends on conditions on the bait grounds and to some extent the biting response of the target tuna species.
Between one and three hauls per vessel per night is usual, with an overall average from the data sources in Table 18 of two hauls per night.
Lewis (1990) has reviewed catch data from the South Pacific Commission Skipjack Programme’s bait fish catches throughout the region. Where bait fishing was
conducted, average catch rates ranged from 34 to 291kghaul−1, with an overall average of 101kg haul−1. Lewis (1990) showed that the highest catch rates (120–291
kghaul−1) occurred predominantly around high islands with extensive areas of sheltered lagoons and embayments. Catch rates from atolls lay mainly in the middle of the
range (57–100 kg haul−1), with the lowest catch rates (34–39kghaul−1) around high islands with limited lagoon area. Little or no bait fishing was conducted at Pitcairn,
Niue, Nauru, Tokelau and Norfolk Islands. Apart from Tokelau, the other four locations are small exposed high islands with limited or no fringing reefs and no lagoon
area. Bait lights were hung from the survey vessel to see if there were any indications of suitable small pelagic species and in each instance the results suggested limited
stocks of bait fish.
Catch rates of many small pelagic fishes in the South Pacific are strongly seasonal. This seasonality becomes more pronounced at higher latitudes where there are
temperature differences of 8 to 10°C between winter and summer. The most consistent observations on fisheries for small pelagic species come from the various Pacific
island bait fisheries. Ysabel Passage (Fig. 2, 118), in New Ireland Province, northern PNG, lies just below the equator at 2°30’S and average monthly temperatures in
this bait ground range from 29.3 to 30.5°C. However, catch rates are strongly seasonal, with highest CPUEs occurring between June to September (Dalzell & Lewis
1988). A similar observation was also noted for catches from another bait ground, Cape Lambert (Fig. 2, 13), lying about 4°S on the north coast of East New Britain,
where CPUEs were highest between June and October (Dalzell & Lewis 1988). Analyses of catch rates for individual species at the Ysabel Passage fishery showed
that the CPUES for the two anchovies, Encrasicholina heteroloba and E. devisi peaked markedly during July and October. Conversely, the CPUE for the sprat,
Spratelloides gracilis, peaked in September. The two production peaks for the anchovies are synchronous with the two periods of peak reproductive activity (Dalzell
1987). There was no apparent relationship between reproductive activity of the sprat and the production peak in September.
Based on data presented by Evans & Nichols (1985), the seasonality of the Solomon Islands bait fishery (8°S) was not so marked, but a peak in CPUE was noted
during June, and this coincides with a peak in spawning activity for the stolephorid anchovies. The Palau bait fishery (7°N), like those in PNG, the Solomon Islands and
New Caledonia was predomi
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nantly a fishery for stolephorid anchovies. Muller (1976) states that over a 10yr period CPUES peaked in MayJuly and NovemberDecember, as a result of an influx
of recruits. The Kiribati bait fishery is composed of catches at four atolls lying between 1 and 3°N of the equator and is dependent mainly on catches of sprats, sardines
and herrings (Table 16). Data presented by Rawlinson et al. (1992) suggest that there are peaks in CPUE during February and between August and September.
Rawlinson et al. found no evidence to link these production peaks with peaks in reproductive activity. Catch rates in the Fijian bait fishery (17°S) are distinctly
seasonal, with a peak in April and May, declining through June and August to a low during October and November (Sharma & Adams 1990). Sharma & Adams
report that there is a strong correlation between surface water temperature and CPUE in the Fijian bait fishery. A similar trend was noted in the bait fish catches from
New Caledonia (22°S), with a similar peak in March and April, and a low CPUE between August to November (Conand 1988).
As all fisheries for live bait in the South Pacific have relied on light aggregation prior to capture, the other major influence on these bait fisheries is moon phase. The
attractive power of the submersible lamps is diminished at the full moon period when the amount of ambient light is greatest. Kearney (1977) has shown that bait catch
rates in PNG were lowest during periods of full moon, while the last quarter catch rates are consistently lower than for new and first quarter catch rates. The catch rates
at the full moon are usually so poor that it is common practice for the tuna vessels to take a 2 or 3day break over this period.
Seasonality of catches of other small pelagic fishes is not as detailed as for the poleandline baitfisheries. Liftnet catches of round scads, sardines and mackerels
peaked between September and November in Chuuk Lagoon, while bigeye scads showed two peaks during April and August (Anon. 1992c). Purseseine catches of
round scads were greatest in April in Vava’u, Tonga but declined thereafter while sardines and goldspot herring increased in dominance in the catch (King 1992b).
There are some indications that there may be an inverse relationship between the abundance of round scads and sardines at Vava’u. Catch rates of big eye scads from
beach seining between December and June showed a clear peak during February in northern PNG with a steady decline thereafter until schools had disappeared from
the shore areas by July. Amesbury et al. (1986) state that bigeye scads can be caught year round beyond the reef at Guam by nighttime jigging, but that there is a
seasonal run of juveniles between August to November that enter bays and harbours where they can be caught by gill nets, handlines and cast nets.

Because of their importance as a supportive fishery for poleandline tuna fishing, bait fisheries in the Pacific have been studied in some detail. Furthermore, as these are
inshore fisheries, bait is caught in lagoons and bays used as fishing grounds by local people. Although the fishers are usually targeting larger reef and lagoon species,
conflicts have arisen over access rights to traditional fishing areas and over what people perceive as interactions between bait fisheries and subsistence fisheries.
A number of studies have been conducted on the various component species of the different bait fisheries in the South Pacific. The most intensively studied of these
various bait species is the Hawaiian anchovy or nehu (Encrasicholina purpurea). Studies of this species include egg and larval production, food and feeding behaviour
and population biology, as well as early investigations into the fisheries biology (Nakamura 1970). Later studies included reproduction and fecundity (Leary et al.
1975), tagging with microtags (Leary &
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Murphy 1975), age and growth from otolith microstructure (Struhsaker & Uchiyama 1976) and further investigations into fisheries dynamics and population biology
(Weatherall 1977). Recent studies include further investigations into the reproductive biology of nehu such as fecundity and spawning frequency (Clarke 1987, 1989)
and estimation of biomass using the egg production method (Somerton 1990, Clarke 1992).
Investigations into the fisheries biology of stolephorid anchovies and other species used for bait fish have been conducted in Palau (Muller 1976), PNG (Dalzell
1990a), New Caledonia (Conand 1988), the Solomon Islands (Evans & Nichols 1985, Blaber & Copland 1990, Blaber et al. 1993), Kiribati (Rawlinson et al. 1992)
and Fiji (Sharma & Adams 1990, Blaber et al. 1993). These investigations have concentrated on the following species: Encrasicholina heteroloba, E. devisi, E.
punctifer, Spratelloides gracilis, S. delicatulus, S. lewisi, Amblygaster sirm and Herklotsichthys quadrimaculatus. Other species that occur regularly in bait hauls
and have been studied in some detail include the hardyheads, Hypoatherina ovalau and Atherinomorus lacunosus; anchovies, Stolephorus indicus, S. insularis and
Thryssa balaema; sardine, Amblygaster clupeoides; round herrings, Dussumieria spp., and ponyfishes, Leiognathus bindus and Gazza minuta (Conand 1988).
Apogonids or cardinalfishes have received some attention as they can form a significant portion of the bait catch. Species include Archamia zosterophora in the
Solomon Islands (Milton et al. 1990) and Rhabdamia gracilis and R. cypselurus in Fiji (Milton et al. 1993). Hoedt (1984, 1990) has studied the biology of anchovies
in northeastern Australia, including Stolephorus nelsoni, S. carpenteriae, S. insularis, Encrasicholina devisi, Scutengraulis hamiltoni and Thryssa setirostris.
Conand’s (1988) study of bait fishes in New Caledonia also included some of the larger small pelagic species such as the bigeye scad, Selar crumenophthalmus,
Indian mackerel, Rastrelliger kanagurta, and the round scad, Decapterus russelli. These species, though important in subsistence and small commercial fisheries,
have largely been ignored in the region, apart from Tonga, where the age, growth and mortality of D. macrosoma and D. macarellus were studied from samples taken
with an experimental purse seine (King 1992b). Also included in the Tonga study were the herring, Herklotsichthys quadrimaculatus, and the sardine, Amblygaster
sirm. The biology and fisheries for bigeye scads in Hawaii have been studied by Kawamoto (1973) and Kazama (1977), while the biology and fishery for the round
scad Decapterus macarellus were studied by Yamaguchi (1953).
The studies of the small pelagic species targeted for tuna bait in the South Pacific have covered most aspects of the life histories of the dominant species. The short
life spans of tropical small pelagic fishes have meant that they are relatively easy to age using otolith microstructure and to estimate growth parameters from otoliths,
length frequencies or a combination of both. Based on such studies, Conand (1988) and later Lewis (1990) proposed that tropical small pelagic fishes could be
separated into two groups based on life history parameters.
Type Species with a short life cycle (less than 1 year), which are relatively small in size (7–10 cm max), grow rapidly, attaining sexual maturity in 3–4 months, spawn
1. over an extended period and have batch fecundities of 500–1500 oocytes per gramme of fish. These include the stolephorid anchovies (Encrasicholina
heteroloba, E. devisi, E. punctifer), sprats (Spratelloides gracilis, S. delicatulus, S. lewisi) and silversides (Hypoatherina ovalau), all of which are important
tuna bait fish species.
Type Species with an annual life cycle (but with some individuals surviving to 2 years of age), which are larger in size (10–24cm max), attain sexual maturity towards
2. the end of the first year, spawn on a restricted seasonal basis and have batch fecundities
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in the order of 300 to 500 oocytes per gramme of fish. These include the herrings and sardines (Herklotsichthys spp., Amblygaster spp., Sardinella spp.), the larger
anchovies (Thryssa spp., Stolephorus spp.) and the sharpnosed sprats (Dussumieria spp.).
Dalzell (1993b) considered that a third category was required to include the larger small pelagic species.
Type Species that have life spans of between 2 and 5yr and that attain sizes of 20 to 35cm maximum size. These species also exhibit restricted spawning seasons and
3. have relatively low batch fecundities (400 to 600 oocytes per gramme of fish), although flying fish batch fecundities are much lower (50 to 100 oocytes per
gramme of fish) in common with Type 2 species. Included in this group are the round scads (Decapterus spp.), the bigeye scads (Selar spp.), the small mackerels
(Rastrelliger spp.) and the flying fishes and halfbeaks (Exocoetidae and Hemiramphidae)
As might be expected from the foregoing, the short life spans (0.4 to 4yr) of these tropical small pelagic fishes give rise to very high mortality rates. According to
Gulland (1983), where mortality rates are high it will pay to fish such stocks relatively hard and with a low size at first capture since many fish will die before completing
much of their growth.
Not surprisingly, the most consistent sources of data on small pelagic fisheries in the South Pacific are from the tuna bait fisheries in PNG, the Solomon Islands,
Palau, Kiribati and Fiji. The simplest approach to managing these fisheries was to attempt to fit simple SchaeferFox surplus production models to catch and effort
data. With the exception of Palau, the relationships between catch and effort for the entire bait fish production in each country were linear and there was little evidence
of curvature. The lack of pronounced curvature in the catcheffort relationship for the PNG, Solomons, Kiribati and Fijian bait catches may be due to the dynamics of
these poleandline fisheries. When catches in a particular bait ground decline, either through localized overfishing or through environmental effects, the poleandline
fleet will usually move to another bait ground. Furthermore, although individual species within a bait catch might decline during a fishing season, there is usually an
increase in abundance of one or more of the dominant species in the catch to compensate for the decline.
Muller (1976) fitted both Schaefer and Fox curves to the data from Palau to obtain an estimate of the MSY and optimum fishing effort. Muller’s estimate of MSY
was 80000 buckets, or 160t, 90% of which would be Encrasicholina heteroloba. For a bait ground of 300km2 this represents a total yield of 0.53tkm−2 or
0.48tkm−2 of E. heteroloba. Actual yields from the Palau bait fishery ranged from 0.29 to 0.67tkm−2 (mean=0.44tkm−2). Dalzell (1984a, 1990a) showed that the
relationship between catch and fishing effort for stolephorid anchovies from the Ysabel Passage and Cape Lambert bait fisheries did not conform to the linear model for
total bait catch. Dalzell (1984a) fitted simple Schaefer surplus production models to the data for catches of E. heteroloba and E. devisi. Estimates of MSY were
initially made with catch data from the Ysabel Passage, but were expanded to incorporate the smaller data set from Cape Lambert. Catch and fishing effort from each
bait ground were standardized between the two bait grounds by expressing them on a per unit area basis. An analysis of the E. heteroloba data alone gave a predicted
MSY of 0.44tkm−2yr−1, similar to 0.48tkm−2yr−1 for the same species in Palau (Muller 1976). Actual yields ranged between 0.2 to 1.2tkm−2yr−1
(mean=0.49tkm−2yr−1) in PNG. The MSY for combined catches of E. heteroloba and E. devisi from Ysabel Passage and Cape Lambert was about 0.65tkm−2yr−1.
The combined actual yields of both anchovies ranged from 0.25 to 1.60tkm−2yr−1 with a mean of 0.75tkm−2yr−1.
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Nichols & Rawlinson (1990) and Tiroba (1993) have analyzed catch and effort data from individual bait grounds in the Solomon Islands bait fishery and showed that
simple Schaefer production curves will fit the data. Nichols & Rawlinson (1990) fitted a Schaefer curve to catcheffort data from 1981 to 1989 for the Munda (Fig. 2,
63) bait ground. Tiroba (1993) repeated this analysis but included additional information on catch and effort from 1990–92. Tiroba was also able to fit Schaefer curves
to data for bait grounds at Mbili (Fig. 2, 58) and Thousandships Bay (Fig. 2, 99). The MSYs for these bait fisheries ranged from 357 to 535tyr−1. Information on the
areas of these bait grounds was not available. Dalzell & Lewis (1989), however, quote a figure for the whole of the Solomon Islands of 806km2 bait fish grounds. The
total bait catch in the Solomon Islands varied from 1800 to 2500t between 1984 and 1988 (Nichols & Rawlinson 1990), suggesting yields of 2.2 to 3.0tkm−2yr−1 with
a mean of 2.7tkm−2yr−1. In terms of stolephorid anchovies only, this represents yields of between 1.6 and 2.2tkm−2yr−1. The total average yield from the two main
PNG bait grounds, Cape Lambert and Ysabel Passage, was 1.1 tkm−2yr−1, less than half the average total yield from the Solomon Islands.
The analyses in the previous paragraphs do not take into account the effects of the environment on the productivity of small pelagic stocks and ascribe changes in
abundance directly to the variations in fishing effort and thus fishing mortality. There is, however, some evidence to suggest that the production of tropical clupeoids, and
possibly other small pelagic fishes, is strongly influenced by environmental effects, particularly wind and rainfall. Dalzell (1984b) investigated the effects of rainfall on
catches of the anchovies E. devisi and E. heteroloba in PNG and suggested yields of these species takes the form of a parabolic function with respect to rainfall, with
optimum rainfall for both species of approximately 3000 mmyr−1. Muller (1976) has indicated that rainfall enhances recruitment of E. heteroloba at Palau. Thus
recruitment, and hence catch rates, of E. heteroloba might be expected to decline during the years that are drier than average.
Other investigations of the effects of rainfall on small pelagic species used as bait fishes have been made in Fiji (Ellway & Kearney 1981) and Kiribati (Ianelli 1992).
Ellway & Kearney (1981) suggested that rainfall did not markedly affect bait fish catches in Fijian waters. The authors, however, did not investigate the effect of rainfall
on individual catch components; rather they used the catch data for all species combined. There was also no significant correlation between rainfall and total catch in the
Kiribati bait fishery, although the scatter of points of catch rate versus rainfall presented by Ianelli (1992) suggest an initial increase in catch rates as rainfall increases,
but with declining catch rates at the highest levels of precipitation. Ianelli (1992) did find, however, a significant positive correlation between catch rate of Amblygaster
sirm and Spratelloides delicatulus and rainfall. Dalzell (1984b) found no correlation between catch rates of the congener S. gracilis from the northern PNG bait
fisheries and rainfall.
The analyses of small pelagic catches presented here are concerned mainly with bait fisheries that catch Encrasicholina spp. and Spratelloides spp. No equivalent
data exist for other small pelagic species such as scads, mackerels, flying fishes and halfbeaks. Unfortunately, the data for making these assessments are presently not
available for most locations. The standing stock biomass per km2 of small pelagic species has been estimated by Petit & Le Philippe (1983) for bays and lagoons
around New Caledonia to be between 0.04 and 1.84tkm−2, with a weighted mean of 0.465tkm−2, consisting primarily of sardines, anchovies and sprats. Estimates of
mean standing stock of small pelagic species, primarily anchovies and sprats, for the Ysabel Passage and Cape Lambert bait grounds of PNG (Dalzell 1984a) were
0.59 and 0.29tkm−2, with an overall weighted mean of 0.42tkm−2. These lim
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ited data suggest that at least around major high islands in the South Pacific, the biomass densities of small pelagic fishes may be relatively similar.
Small pelagic fishes will continue to be an important component of subsistence fisheries in the South Pacific, particularly where reef and lagoon areas are limited. For
example, Chambers (1984) notes the importance of flying fishes in the diet of people living on the small coral island of Nanumea (Fig. 2, 65), where almost a quarter of
all meals contain flying fishes. The limited size of most islands and the surrounding shelf area mean that the small pelagic resources are also limited. Thus smallscale
commercial enterprises can survive but largescale exploitation of stocks is unlikely because of resource limitation. Information on coastal fisheries in Western Samoa
suggests an overall decline in the amount of fish sold through Apia market. Some of this decline may be due to fish being exported to neighbouring American Samoa
where better prices can be realized; but there is also some indication that some stocks, including bigeye scads, have suffered real declines in abundance, with sales of
less than 1.0tyr−1 through the market in 1990 compared with 12.5t in 1986 (FAO 1990).
The demand for bait fish for poleandline fishing is unlikely to grow and indeed the present trend is one of reduction in this method of fishing for tuna. The last large
scale poleandline fishery is in the Solomon Islands, which at its peak in 1985 contained 36 boats but by 1993 was reduced to a fleet of 27 boats, and is expected to
be reduced even further as poleandline fishing is replaced by purse seining. There is the possibility of using some of the larger pelagic species (scads and sardines) for
longline bait as more countries establish mediumscale longline fisheries for the export of very highvalue tuna. Such supportive bait fisheries have been established in
Southeast Asia but, as stated earlier, attempts to use locally available small pelagic species for longline bait in Fiji were confounded by seasonality of abundance and
the unreliability of supplies. Longline fishing is thus likely to continue to rely on commercially produced bait from Japan despite the cost of purchase and shipping to the
South Pacific.
The smaller species such as the stolephorid anchovies are caught and consumed in considerable quantities in Southeast Asia, where production is now approximately
250,000tyr−1 (FAO 1994). Anchovies are eaten fresh, dried or fermented into fish pastes and sauces (Ruddle 1986). This level of production in Southeast Asia
appears to be sustainable, although fishing pressure is very high and, given the low unit value of these fishes in Asia, the possibilities of exporting to this neighbouring
region are minimal. Such types of small pelagic fish command very low prices in Asia and a fishery for them based, for example, on a dried product is not profitable
under present circumstances, as indicated by the shortlived interest of Asian entrepreneurs entering the Pacific islands region.
Estuarine fisheries
Description
Estuarine fisheries are of importance to only a few countries in the South Pacific region, the majority of islands being small with limited or nonexistent estuarine areas.
However, in terms of population, a relatively large number of people are dependent on estuarine fisheries resources, given that the major estuarine areas are found in
the large island archipelagos of Melanesia. The biggest estuarine area in the South Pacific is the Gulf of Papua. Several
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major river systems drain from the highlands southward into the Gulf of Papua, including the FlyStrickland system (Fig. 2, 25), and the Kikori (Fig. 2, 40) and the
Purari (Fig. 2, 82) Rivers. In addition, several major rivers discharge along the north coast of PNG, namely the Markham (Fig. 2, 55), Ramu (Fig. 2, 85) and Sepik
(Fig. 2, 92) Rivers, creating smaller but still extensive estuarine areas. The other Melanesian islands have smaller river systems and much smaller estuarine areas.
However, in countries like Fiji these estuaries and associated wetlands such as mangrove and Nipa palm forests contain important fisheries resources. In most of the
other Pacific high islands, the land masses are too small to have any significant river systems. Usually, only small streams are found and these may be seasonal in
occurrence. Islands such as Pohnpei and Kosrae in the Federated States of Micronesia experience prolonged and extensive wet periods, but despite the runoff, are
surrounded by well developed fringing reefs and, in the case of Pohnpei, a barrier reef as well.
As described earlier, trawlers fishing for shrimps along the coastal margin and on the coastal shelf of the Gulf of Papua catch predominantly estuarine and soft
bottom associated fishes. Watson (1984) estimates that the annual harvest in this fishery of finfish alone amounts to between 11300 and 17000t (mean 14000).
Elsewhere on the northern PNG coast, limited trawl fishing has been conducted at various sites, either to estimate fisheries resources or as part of investigations into
estuarine fish populations. Coates et al. (1984) conducted a 3month survey of the trawlable grounds at the head of Milne Bay (Fig. 2, 60), which is estuarine in nature
due to the numerous streams and rivers that discharge into this region. Survey trawling has also been conducted off the mouth of the Sepik River and within the Murik
Lakes (Fig. 2, 64), which form part of the Sepik Delta region (Chapau 1991). The estuarine fish and shrimp stocks of the Markham River estuary have been studied in
detail by Quinn & Kojis (1986) using smallscale beam trawling.
Opnai (1984, 1986a) described the fisheries of the northern Gulf of Papua centred around the fisheries centre at Baimuru. As part of a fisheries development
project, gill net fishing was encouraged in the estuarine and fresh waters in proximity to Baimuru (Fig. 2, 7) to improve fisheries production and income opportunities in
an area where there was little agricultural land. Village fishermen set gill nets from dugout canoes and kept fish in ice boxes to be picked up by a collection vessel for
transport to the fisheries center at Baimuru. This fishery produced between 30 and 100tyr−1 (mean=58tyr−1). Similar gill net fisheries are found along most of the length
of the Gulf of Papua, and around the Fly River mouth. One of the principal targets for these fisheries is the catadromous sea bass, barramundi (Lates calcarifer), which
occurs naturally in the South Pacific islands only in PNG, with a range that extends from the Irian Jaya border in the west to the south eastern tip of the Papuan coast
near Samarai.
The subsistence fisheries of the neighbouring Purari Delta have been described by Haines (1978/79). Traditional methods for catching fishes in the Purari include
handlines, traps, spears and bows and arrows. Besides fishes, villagers in the Purari region catch mud crabs and a variety of shrimps including the freshwater
Macrobrachium spp. The finfish catches are dominated by ariid catfishes, barramundi and sharks. Gill nets introduced by fisheries extension officers have gradually
replaced traditional fishing methods and are responsible for generating most of the catch. Most of the fisheries production is for subsistence, with only a small amount
being traded by those villages without suitable agricultural land. Haines estimated that the average catch of crabs and fishes in the Purari villages amounted to about 40
and 80g day−1 respectively.
The barramundi fishery in the Western Province of PNG is divided into two components; the coastal fishery operating between the Fly River mouth and the mouth of
the Binaturi
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River (Fig. 2, 10) to the west of Daru (Fig. 2, 18), and the inland fishery based in the Fly River and Lake Murray (Fig. 2, 44) (Opnai & Tenakanai 1987). The two
fisheries are based on different aspects of barramundi life history. Briefly, this species spawns in marine coastal waters and larval/juvenile stages migrate into coastal
swamps during the wet season. As the wet season finishes and the floodwaters recede, the youngoftheyear fish move upstream to the upper reaches of the river
where they remain for 3 to 4 yr. After this period of residency upstream the fish become sexually mature males and migrate to the tidal waters downstream to
participate in spawning and to ultimately change into females between the ages of 6 to 8 yr. Following spawning, the females disperse along the coast until the next
spawning season. Catches within the Lake Murray area are taken throughout the year due to the presence of prespawning year classes that do not migrate to the
coast. The coastal fishery experiences low catches of barramundi throughout much of the year, except between October and February when adult fishes migrate to the
coastal areas to spawn.
Quinn (1984) described the smallscale subsistence and commercial fisheries of the Markham River estuary on the north coast of PNG. Fishers from the villages on
the estuary catch fishes, shrimps and crabs for food and for sale in the nearby town of Lae. Fishermen use bows and arrows, poleandline and nets to catch fish. Bow
and arrows are used from canoes to catch jewfish, trevallies and eels. Fishermen catch shrimps to be used as bait for poleandline fishing for snappers at the mouth of
the Markham River. Shrimps are usually caught by a scoop net that is pushed through the water for about 2 to 4 min. Other species caught include juvenile trevallies,
and mullet. Beach seines and gill nets are also used by fishers in the Markham estuary to catch trevallies, mullets and crabs. Small ovalshaped nets are also set in
among the mangrove roots to catch crabs and gudgeons.
Close to the mouth of the Sepik River is a series of coastal lagoons that connect with the Sepik and open out into the Pacific Ocean, known as the Murik Lakes. The
Murik Lakes system covers about 90km2 and the associated mangroves, about 185km2. The fisheries of these lagoons have been studied by Chapau (1991) and are
among the most productive coastal fisheries along the north coast of PNG. Production from the Murik Lakes accounts for twothirds of the commercial fish landings to
the nearby urban center of Wewak. The commonest fishing methods employed to catch finfish in the Murik Lakes are hand lining and gill netting from canoes. Besides
finfish, the villagers of the Murik Lakes also catch mud crabs in the mangroves and harvest various oysters and clams. A similar area of sheltered coastal estuarine
water, Sissano Lagoon (Fig. 2, 93), is found further west along the northern coast of PNG. Sissano Lagoon covers an area of about 11km2 and is formed by the
drainage of a large mangrove swamp that is restricted by coastal sand bars. As with the Murik Lakes, Sissano Lagoon is one of the few areas of sheltered coastal
water on the northern PNG coast where fishermen can fish all year round. The fisheries of Sissano Lagoon are broadly similar to those of the Murik Lakes, with finfish
catches being made predominantly by gill nets and handlines (Ulaiwai 1992a,b).
Little information is available on estuarine fisheries elsewhere in the South Pacific. Lauvi Lagoon (Fig. 2, 47) on the south coast of Guadalcanal (Fig. 2, 31) in the
Solomon Islands is a coastal lagoon of 3km2 in area, similar to the Murik Lakes and Sissano Lagoon in PNG. Unlike the PNG lagoons, Lauvi Lagoon is mostly fresh
water for most of the year, being completely enclosed by sand bars that are broken only during the rainy season (JulyAugust). Gray (1974) describes the fish fauna of
Lauvi Lagoon and from other fresh and brackish waters on the Solomon Islands, while Hinds (1971) provides some details of fish catches made there with gill nets.
Batty (1987) gives some details of eel catches made with fyke nets on Guadalcanal, although these were predominantly in fresh rather than brackish waters.
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The fresh water and estuarine fisheries of Fiji’s main island, Viti Levu, have been described by Lewis & Pring (1986) and more recently by Rawlinson et al. (1994).
These authors list about 35 fishes that might be considered euryhaline or brackish water species. These include such species as the introduced exotic freshwater cichlid
or tilapia, Oreochromis mossambicus, and the jungle perch, Kuhlia rupestris, which is a catadromous species like the barramundi in that adults swim down to the sea
to spawn. This species is widely distributed in the Pacific and can be found on most high islands where there are permanent streams or rivers. Lewis & Pring noted that
it was difficult to assess just what quantity of fish are produced from estuarine waters in Fiji, as some species are taken in both brackish and coralline environments. The
total annual commercial catch of estuarine species listed by Lewis & Pring is about 1100t, of which, they suggest, only 10% is taken in brackish waters. Rawlinson et
al. (1994) state that the most frequently reported fish species caught in rivers and estuaries were tilapia, eels, jungle perch and the snapper, Lutjanus
argentimaculatus.
David & Cillauren (1989) report no estuarine fishing in Vanuatu, but there is a small freshwater fish catch of about 50tyr−1. Exploitation of estuarine resources in
New Caledonia appear to be mainly limited to mud crabs (see p. 488). Fishermen in Noumea will occasionally catch tilapia and mullet in tidal creeks and inlets using
cast nets and gill nets (Dalzell, pers. obs.). The fish fauna of mangrove systems and estuaries in proximity to extensive reef systems in the New Caledonia lagoon have
been studied by the French scientific organization, ORSTOM. Wantiez (1993) has conducted a series of experimental trawl fishing surveys of areas of the New
Caledonia coast with two types of trawl net, one of which was designed to target shrimps. Thollot (1992) has sampled populations of estuarine fishes in mangrove
lagoons at several sites in New Caledonia, using principally gill nets but also with trammel nets and barrier nets.
As this is a review of coastal fisheries, we have touched only obliquely on freshwater fisheries production. As with estuarine fisheries production, freshwater fisheries
are of limited importance in most locations in the South Pacific. In all but the largest high islands, streams and rivers in the South Pacific are short and in some cases
seasonal, depending on rainfall regime. As is clear from the foregoing the largest rivers are found in the Melanesian islands, and the largest rivers in the South Pacific
flow through the mainland of PNG. The freshwater fauna of the South Pacific is very impoverished, with much of it secondarily derived from marine fishes. It is
therefore not surprising that the major freshwater fish stocks of PNG are all introduced species, tilapia (Oreochromis mossambicus), common carp (Cyprinus carpio)
and the rainbow trout (Oncorhynchus mykiss). The scale of freshwater fish production from PNG is unknown, but for the Sepik River flood plain alone, Coates
(1985) estimates a total annual yield of between 3000 and 5000t.
Catch composition
Descriptions of the mangrove and estuarine fish fauna are given for PNG by Kailola & Wilson (1978), Collette (1983), and Quinn & Kojis (1986), for the Solomon
Islands by Gray (1974) and Blaber & Milton (1990) and for New Caledonia by Thollot (1992), Kulbicki & Wantiez (1990) and Wantiez (1993). The catch
compositions from fishing in estuarine locations in PNG, New Caledonia and Fiji are shown in Table 19. Hinds (1971) does not give the relative amounts of each
species caught in Lauvi Lagoon, Solomon Islands, by gill net fishing but states that the catches included milkfish (Chanos chanos), trevally (Caranx ignobilis), mullet
(Mugilidae), tarpon (Megalops cyprinoides) and mangrove jack (Lutjanus argentimaculatus). A feature of the estuarine and softbottom catches in the South Pacific
is
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the extreme variability in the relative contributions by the different fish families. There are large numbers of species captured in other coastal demersal fisheries such as
from reef and deepslope stocks, however the majority of the catch tends to be formed from the same combination of families. This does not appear to be the case
with estuarine fisheries, although this may be partly because of the variety of gears being deployed, which range from simple handlines and spears to large scale trawls,
and the different types of location, which include small localized estuaries, coastal estuarine lagoons and a large estuarineinfluenced continental shelf.
Trawl catches on the coastal shelf of the Gulf of Papua are dominated by jewfishes, clupeoids (anchovies and sardines) and pony fishes. Kailola & Wilson (1978)
have compared the fish fauna of the eastern and western sectors of the Gulf of Papua. They showed that in the east the trawl finfish catches are dominated by
ponyfishes, bream (Nemipteridae), grunters (Theraponidae), snappers, jacks and silver biddies (Gerridae), while in the west there are comparatively higher percentages
of catfishes (Ariidae), jewfishes, threadfin salmon (Polynemidae), anchovies and bombay ducks (Harpontidae). Kailola & Wilson state that in the west, muddy water
from the deltas of the northern Gulf create conditions of low visibility, favouring those fishes that show adaptations typical of muddy water forms, such as sensory
barbels, projecting snout and adipose eyelids. To the east, where the waters are generally less turbid, those species associated with shallow coastal waters and sandy
beaches are found.
Both the catches from Milne Bay (Coates et al. 1984) made with an otter trawl, and from the Markham River estuary (Quinn & Kojis 1986) made by smallscale
beam trawling from an outboard powered dinghy, contained predominantly ponyfishes. However, the Milne Bay catch also contained significant quantities of hairtails
(Trichiuridae) and javelinfishes (Pomadasyidae), while the balance of the Markham River catch were the toadfishes (Tetraodontidae) and snappers. Quinn & Kojis
(1985) also found that ponyfishes formed over threequarters of small beam trawl catches at a small estuary, Mis Inlet (Fig. 2, 61), adjacent to a major reef system
near Madang (Fig. 2, 50), northern PNG. A similar small scale trawl deployed in the lagoon of Tongatapu, the main island of Tonga, caught a range of estuarine and
coral reef associated species. The catch was dominated, however, by snappers, ponyfishes and silver biddies (Braley 1976).
Commercial gill net catches around Baimuru in the northern Gulf of Papua (Anon. 1985) comprised mainly barramundi, with threadfin salmon, catfishes and jewfishes
making up most of the remainder of the catch. Subsistence catches in the neighbouring Purari Delta comprised mainly catfishes, barramundi, threadfin salmon and
sharks (Haines 1978/79). Subsistence catches with gill nets along the north coast of PNG were dominated by jacks, snappers, milkfishes and threadfin salmon at the
Murik Lakes (Chapau 1991), and sardines, mullet, silver biddies (Gerridae) and ponyfishes at Sissano Lagoon (Ulaiwi 1992b). Handline catches at the Murik Lakes
comprised mainly javelinfishes, jacks and snappers (Chapau 1991).
Experimental catches by gill nets in mangrove lagoons in New Caledonia comprised mainly mullet, rabbitfishes, snappers and sardines (Thollot 1992). Catches made
in similar locations with a fish corral or barrier net were dominated, in terms of weight, by groupers, grunts (Haemulidae), silversides and rabbitfishes. Due to the small
mesh sizes employed, however, the silversides formed about twothirds of the fish corral catch by numbers. Experimental trawl catches on softbottoms adjacent to
mangrove estuaries in New Caledonia were dominated by the ponyfishes, goatfishes and emperors (Kulbicki & Wantiez 1990).
The composition of the estuarine catch from Fiji is based on those species defined by
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Table 19 Percent catch composition of estuarine and soft bottom fisheries in the South Pacific region.
Location and fishery

Family Gulf of Milne Baimuru gill Purari Delta Markham Murik Murik Sissano New New New Fiji gill
Papua Bay net catch gill nets and River mouth Lakes Lakes Lagoon gill Caledonia Caledonia Caledonia nets and
trawl trawl other gears other gears trawl gill net handline net gill net fishcoral trawl other
gears
Sciaenidae 22 0.8 9.4
Engraulidae 15 3
Leiognathidae 10.3 42.8 66.9 12.1 1.6 3.7 42 2.7
Trichiuridae 7.8 13.3 1.1
Harpontidae 6.8
Clupeidae 5.7 19.8 8.6 1.4
Pomadasyidae 4.8 9.8 11.7 48
Mullidae 4.6 5.9 1.9 14 2.5
Ariidae 3.6 1 8.1 23.5
Theraponidae 2.9
Polynemidae 1.8 2.3 13.8 12 2.7
Sauridae 6.7
Carangidae 2.6 38.8 12.9 8.3 1.1 37.7
Sphyraenidae 0.6 4.8 3 1.9
Centropomidae 59.7 15.3
Eleotridae 3
Lutjanidae 3.9 16.5 14.1 2.2 8.8 1.9 8.2
Chanidae 13
Mugilidae 5.3 27.9 19.5 L2 43.2
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Location and fishery

Family Gulf of Milne Baimuru gill Purari Delta Markham Murik Murik Sissano New New New Fiji gill
Papua Bay net catch gill nets and River mouth Lakes Lakes Lagoon gill Caledonia Caledonia Caledonia nets and
trawl trawl other gears other gears trawl gill net handline net gill net fishcoral trawl other
gears
Scatophagidae 4 3.8
Haemulidae 1 7.6 1.9 8.7 17.6
Serranidae 5.1 3.3 25.6
Gerridae 1.9 13.3 5.9 4.5 2.2
Lethrinidae 1.7 1 2.7 3.6 9.8
Tachysuridae 0.9
Megalopidae 0.7
Siganidae 1.4 12.5 11.3
Scaridae 3.2
Sparidae 3.9
Plotosidae 2.1
Siliganidae 2.1 1
Muraenidae 1.4 2.6
Atherinidae 14.3
Apogonidae 2
Monodactylidae 2
Platycephalidae 1.5
Synodontidae 3.7
Nemipteridae 1.3
Tetraodontidae 14.5 1.1
Bothidae
Hemiramphidae 2.8
Lactaridae 1.1 3.1
Sharks 4.7 24.6
Others 14.7 14.8 4.3 18.6 9.9 10.7 2.7 1.4 9/7 3.9 25.9 4.6
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Lewis & Pring (1986) and contemporary catch statistics (Anon. 1994b). Fishes are caught with gill nets and a variety of other gears such as fish corrals, handlines and
spears. Based on the available information, mullet, jacks and snappers comprise most of the commercial estuarine and brackish water catch in Fiji. Rawlinson et al.
(1994) report that the most important subsistence fish catches from rivers and estuaries include tilapia (Oreochromia mossambicus), eel (Anguilla obscura),
mangrove snapper (Lutjanus argentimaculatus), jungle perch (Kuhlia rupestris) and assorted jacks (Carangidae).
Catch rates
Watson (1984) reported that catch rates of finfish in the Gulf of Papua shrimp fishery ranged from 83 kgh−1 to 690kgh−1 with an average of 209kgh−1. Survey fishing
in Milne Bay with similar trawl gear produced catch rates ranging from 60 to 550kgh−1 with a mean of 152kgh−1, comparable to the Gulf of Papua. Commercial trawl
surveys have also been conducted in the coastal waters adjacent to the Murik Lakes (Chapau 1991) although no details of catch composition, other than shrimp weight
and fish weight, are available. Catch rates of fish in three separate surveys ranged from 12 to 600kgh−1 with an overall mean of 92 kgh−1. From their experimental trawl
fishing survey of New Caledonia’s estuarine and soft bottom areas, Kulbicki & Wantiez (1990) reported an average catch rate of 51 kgh−1. The average catch rate of
finfish and shrimps from trawling in the Markham River estuary with a small 3m beam trawl, amounted to 3.4kgh−1 or 3.1kgh−1 of finfish only. Quinn & Kojis (1985)
present no weight data for similar beam trawl catches from the Mis Inlet, but state that average catch rates by numbers were not significantly different from those in the
Markham River mouth. The same type of trawl gear was deployed within the Murik Lakes and in the immediate coastal waters (Chapau 1991) and experienced finfish
catch rates of 0.0–0.53kgh−1 (mean=0.12kgh−1) and 0.15–4.24kgh−1 (mean=1.60kgh−1) respectively.
Catch rates for smallscale commercial gill netting in the estuaries of the northern Gulf of Papua, as described by Opnai (1986a), ranged from 3.6 to 77.5kgset−1,
with an average of 14.5 kg set−1 (Anon. 1985). Nets were usually set in the evening and hauled in the morning. Haines (1978/79) does not present specific information
on catch rates for the subsistence fisheries of the Purari Delta, but based on his data it is possible to make a rough estimate of the average gill net catch rate of
3.3kgnet−1day−1. Gill net catch rates in the Murik Lakes ranged from 3.5 to 6.5kgnet−1day−1, with an overall mean of 5.3kgnet−1 day−1. Handline catch rates in the
same location amounted to between 1.3 and 4.9kgday−1, with a mean of 3.0kgday−1. Catch rates for set gill net fishing in Sissano Lagoon ranged from 0.18 to
0.75kgman−1h−1, with a mean of 0.45kgman−1h−1. This was equivalent to an average catch rate of 3.1kgnet−1 day−1. Catch rates for surround gill netting, where the
net does not fish passively but fish are driven into it by fishermen, ranged from 0.02 to 4.23 kg man−1 h−1, (mean=0.59kgman−1h−1), equivalent to an average catch
rate of 7.3kgnet−1day−1.
Thollot (1992) does not give explicit details of catch rates from gill net fishing or fishing with a barrier net and fish corral in New Caledonia. From his data, however,
it is possible to estimate average catch per set and per hour. Altogether a total of 115 sets of 6 duration were made, generating a total catch of 1114kg, or an average
catch rate of 9.7kgset−1 of 1.6kgh−1. Similarly, a total of 69.1 kg of fish were caught by the barrier net and fish corral, from 24 sets of 12h duration. This gives an
average catch rate of 2.9kgset−1 or 0.24kgh−1.
Page 471

Estuarine resources in the South Pacific are mainly limited to PNG, with limited importance in the other Melanesian islands. Most of the studies on estuarine fish biology
have been conducted in PNG. Barramundi is the most commercially important of the finfish caught along the southwest and central Papuan coast and the biology of this
species has been studied in some detail in PNG, in Australia and Southeast Asia. Reynolds (1978) gave a detailed account of the biology and population dynamics of
barramundi from observations conducted on populations from the Fly River system and the coastal areas around southwest Papua. Reynolds estimated age and
growth, mortality rates and yield per recruit of barramundi and suggested that the levels of commercial and subsistence catch prevalent at that time (early 1970s) were
about optimum as suggested by the yield per recruit analysis.
Further research on barramundi in PNG has been conducted by Moore (1979), who confirmed an earlier observation by Reynolds (1978) that barramundi were
protandrous hermaphrodites. Moore (1982) has also described in detail the reproductive biology and early life history of barramundi, while Reynolds & Moore (1982)
and Moore & Reynolds (1982) have produced further studies on age and growth of barramundi and discussed the results of tagging experiments designed to determine
the migration routes of this species. Commercial landings data between 1971 and 1991 from Daru, the main urban centre of southwest Papua, indicate that barramundi
production was 200–400tyr−1 between 1971 and 1981, but then steadily declined thereafter with annual production in the range 10 to 60tyr−1. The exact cause of this
decline is unknown, and a number of factors have been implicated including pollution from mining tailings disposal from a copper mine upstream, overfishing of the
barramundi stock, or simply a decline in the infrastructural support for fisheries development.
Coates (1987) gives an account of some aspects of biology of the oxeye herring or tarpon, Megalops cyprinoides, from the Sepik River. Coates (1988) has also
studied the fecundity of ariid catfishes on the Sepik River. Coates noted that the very low fecundities and breeding behaviour of these species suggest that recruitment is
likely to be highly densitydependent and the stocks vulnerable to increased mortality, i.e. from fishing. In Fiji, the biology of the catadromous jungle perch, Kuhlia
rupestris, has been studied in some detail as this species has importance for sport fishing and tourism. A synopsis of the results of these studies is given by Lewis &
Hogan (1987).
Few other estuarine fisheries or species have received much attention from fishery biologists in the South Pacific region. Despite the large volume of fish caught each
year in the Gulf of Papua trawl fishery, no biology or stock assessment studies have been conducted, other than Watson’s (1984) initial investigations. Given Watson’s
estimate of average annual finfish production of 14000t for the Gulf of Papua trawl fishery, this amounts to a yield of 1.46tkm−2, based on a total swept area of
9600km2 (Gwyther 1982). Present yields of finfish from the smaller artisanal estuarine fisheries on the northern coast of PNG range from 0.75tkm−2 at the Murik
Lakes (Chapau 1991) to 3.1 tkm−2 at Sissano Lagoon (Ulaiwai 1992b). Information on yields from other estuarine fisheries in the South Pacific are not available. The
studies conducted in New Caledonia on mangrove and estuarine species by Wantiez & Kulbicki (1991), Thollot (1992) and Wantiez (1993) were experimental
observations concerned mainly with community dynamics and changes in population structure in temporal and spatial dimensions. These studies did, however, permit
estimation of biomass of fishes on softbottoms in two locations in New Caledonia that ranged from 0.83 and 4.31tkm−2.
Page 472
The region’s largest catches of estuarine and softbottom species are from the Gulf of Papua shrimp fishery. Watson (1984) estimates that the annual harvest of finfish is
between 11300 and 17200t (mean=14000t) almost all of which is dumped back in the sea. It is uneconomical for the fishing companies to give over freezer space to
the fish bycatch which could only be sold in PNG for a fraction of the shrimp value. The size of the bycatch and the demand for fresh fish in PNG periodically
stimulates demands from politicians that more of the finfish catch from the trawlers is recovered for human consumption. However, it is likely that the economics of the
fishery will continue to dictate that the bycatch will be discarded.
Estuarine and brackish water fisheries will continue to form important components of subsistence diets, particularly along the central and southwestern coast of PNG
and the rural areas of Viti Levu in Fiji. Lewis & Pring (1986) note that brackish water species are of less value than reef and pelagic fishes in Fiji, however estuarine
fisheries remain the most important target species for the rural population of Viti Levu (Rawlinson et al. 1994). Barramundi is probably the single most economically
valuable estuarine species in the region but some form of management initiative is required to see if the decline in production is caused by the collapse of the stock or
related to other factors such as decline in fishing effort for this species.
In French Polynesia, barramundi has been cultured successfully for domestic markets but this culture was not economically feasible and has since been discontinued
(Anon. 1994c). Aquaculture of barramundi in PNG might offer the potential to replenish overfished populations if this was the reason for the decline in production.
Barramundi and other riverine and estuarine species such as K. rupestris may have other intrinsic value in terms of sports fishing and tourism, as is also the case with
species such as the snappers, Lutjanus goldei and L. johnii, that frequent river mouths and estuaries.
Invertebrate fisheries
Echinoderm fisheries
Description
Seacucumbers (Holothuroidea) and seaurchins (Echinoidea) form part of the subsistence diet of some Pacific islanders and can be easily collected during low water
by picking them off the reef, or free diving in shallow waters. This form of reef fishing is often conducted by women and children during the day while the men are at
work elsewhere or out fishing on the reef or open sea. Seaurchins are harvested for the gonads or roe, which are found adhering to the inside test wall. Sea
cucumbers can be eaten fresh, cooked or pickled in lime juice. Mathews & Oiterong (1991) and Smith (1992a) report that pickled seacucumbers are sold in the
markets of Palau and Pohnpei, respectively, while Zoutendyk (1989) states that species of seacucumber are locally important in the Cook Islands for subsistence
purposes. Fermented seacucumber roe is commonly on sale by the roadside in Western Samoa, and is said to be a tonic, particularly during pregnancy, and the
expressed Cuverian tubules of certain Bohadschia species are used by Palauan youngsters to coat the soles of the feet and protect them while walking on the reef
(Adams et al. 1994). In Fiji, only the sandfish, Holothuria scabra, is commonly eaten as a subsistence food source (Adams 1992a), and in the Solomon Islands it
appears that seacucumbers are not traditionally eaten at all (Adams et al. in press).
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Harvesting seacucumbers for processing into bêchedemer (or trepang) is among the oldest of the commercial fisheries in the South Pacific and dates back to the
early 1800s when traders followed the first explorers into the region (Ward 1972). The major producers of bêchedemer were, and still are, PNG, Solomon Islands,
New Caledonia and Fiji. Current export production from the region ranges between 1500 and 2000t annually of dried product (equivalent to 15–20000tFW).
Production figures for some locations such as Kiribati and Tonga are unavailable, but are minor. The main consumers of bêchedemer are Chinese and the majority of
bêchedemer is exported to Hong Kong, Singapore and Taiwan, where much of the lower grades are reexported to the Republic of China. Small volumes of bêche
demer are also sent to speciality Chinese markets in Canada, USA, New Zealand and the UK.
Methods of fishing for, or collecting, bêchedemer are straightforward and there are few differences between countries in terms of the methods employed. In many
situations, holothurians can be harvested by gleaning at low tide, keeping the collected animals in a dinghy or floating container. Crean (1977) describes a method by
which surface swimmers can catch seacucumbers in deep water. A heavily weighted barbed hook attached to a string can be dropped on the animal to pierce and
snag the tough integument so that it can be retrieved. The seacucumbers are processed soon after capture by cooking the animals, usually removing the viscera, and
sometimes smoking the remaining somatic tissue and integument and then drying. Following this procedure, the animals will lose about 90% of their body weight and
shrink by about 50% or more in length. The processing facilities are usually adjacent to the harvesting areas and do not require much capital investment, each consisting
of simple cauldrons for boiling, drying racks and a smoke house. Such structures are easily constructed so that processors can move on to other harvesting areas once
the current area of operations is exhausted.
Catch composition
Mathews & Oiterong (1991) list several echinoderms caught and used both for subsistence and for sale in commercial markets on Palau, including the seacucumbers
Stichopus variegatus, Actinopyga miliaris, A. echinites, Holothuria scabra, H. verrucosa, and the seaurchins Tripneustes gratilla, Hemicentrotus
pulcherrimus, Strongylocentrus pileolus, Toxopneustes pileolus and Diadema setosum. Smith (1986) reports that echinoderm species, such as the seaurchins
Tripneustes gratilla and Echinometra mathaei, and seacucumbers, such as Stichopus horrens and Holothuria atra, are also part of the subsistence diet on Guam.
Preston (1990b) notes that H. scabra is harvested for food as well as bêchedemer production in Fiji. Approximately 7t of seacucumbers are sold for food through
produce markets and stores each year in Fiji (Anon. 1994b), and the seaurchin Heterocentrotus mammilatus can be seen occasionally for sale in both Fiji and
Tonga. Conand (1989) notes that seacucumbers are eaten raw in Wallis and Futuna and grilled in PNG, but no details of species are given. In the Cook Islands,
Zoutendyk (1989) reports that Actinopyga mauritania, Holothuria leucospilota and H. cinarescens are part of the subsistence diet in Rarotonga. Richards (1993)
also notes that subsistence diets in the Cook Islands include the roe of seaurchins such as Heterocentrotus mammilatus and Tripneustes gratilla.
Although gross statistics on commercial exports of bêchedemer are usually good, making this one of the most easily quantifiable of fisheries in the South Pacific,
this information is not necessarily very useful for management. Over 22 species are now commercially traded in western Melanesia and while these all occupy different
habitats they are usually classified only as “bêchedemer” when exported (Adams 1993b). Preston (1993) lists 17 species that
Page 474
Table 20 Commercially exploited seacucumbers in the South Pacific region.
Scientific name Common name

Actinopyga echinites Deepwater redfish
A. leconora Stonefish
A, mauritania Surf redfish
A. miliaris Blackfish
Holothuria atra Blackfish
H. fuscogilva White teatfish
H, fuscopunctata Elephant’s trunk fish
H. nobilis Black teatfish
H. scabra Sandfish
H. edulis Pinkfish
Stichopus chloronotus Greenfish
S. variegatus Curry fish
Thelenota ananas Prickly redfish
T. anax Amberfish
Bohadschia marmorata Chalkfish
B. argus Leopardfish
B. vitiensis Brown sandfish
are commercially harvested and these are given in Table 20. The species most sought after are the black and white teatfish (Holothuria fuscogilva and H. nobilis), the
sandfish (H. scabra) and the prickly redfish (Thelenota ananas) (Anon. 1994f). However, as demand for bêchedemer has continued to grow, lowvalue species
that were originally not considered worth processing into bêchedemer are now being harvested. Recently, a very lowvalue species, Holothuria atra, has begun to
attract the interest of processors as supplies of the other species diminish.
Unlike finfish fisheries in the Pacific where selectivity by the common gears is relatively broad, harvesting of seacucumbers involves visually searching for and
selecting particular species. The first harvests from unfished islands are likely to consist of the highvalue species such as teatfish or sandfish. As harvesting progresses
and the highvalue species populations are reduced, other less valuable species will then be harvested. Dalzell (1990b) reported that bêchedemer production from the
Carteret Islands of eastern PNG was based mainly on H. nobilis. Similarly a pilot bêchedemer fishery in Tuvalu targeted H. nobilis (Pita 1979), while production of
bêchedemer on Ontong Java (Fig. 2, 73) in the Solomon Islands, was based on a mix of H. nobilis and Actinopyga miliaris (Crean 1977). This latter species (or
species assemblage, see Preston 1993) was the main target of bêchedemer processors in Fiji in the late 1980s (Preston 1990b), where harvest levels rose rapidly in
the same decade from 20–30t of dried product in 1984 to over lOOOt in 1988 (Adams 1992b).
Catch rates
Catch rates in terms of seacucumbers usually refer to the number of individual animals collected by one collector in a unit time period, or pieces man−1h−1. Some
catch or collection rates for various seacucumber species are summarized in Table 21. Shelley’s (1981) estimates do not refer to commercial or subsistence catches,
but to counts made during unit times of 1h in Bootless Bay (Fig. 2, 11) on the South Papuan coast. Mathews & Oiterong (1991) reported that average catch rates for
Actinopyga spp., Holothuria scabra and Stichopus variegatus in
Page 475
Table 21 Catch rates of seacucumbers for bêchedemer processing in the South Pacific region.
Location Species CPUE (pieces h−1) Source

range mean
Palau Actinopyga miliaris and A. echinites 18.5–118.0 68.2 Mathews & Oiterong 1991
Palau Holothuria scabra 42.9–100.0 65.5 Mathews & Oiterong 1991
Palau Stichopus variegatus 48.8–71.4 61.7 Mathews & Oiterong 1991
Palau Holuthuria spp. 18.4–68.0 47.1 Mathews & Oiterong 1991
Ontong Java (Solomon Is) H. nobilis 6.0–19.2 11.1 Crean 1977
Fiji H. fuscogilva 12–20 16 Gentle 1979
New Caledonia Thelenota ananas 5–110 42 Conand 1990
New Caledonia H. nobilis 3–190 31 Conand 1990
New Caledonia A. echinites 90–150 118 Conand 1990
Papua New Guinea T. ananas 1.5–70 10 Shelley 1981
Papua New Guinea H. nobilis 1.5–190 15.1 Shelley 1981
Papua New Guinea H. fuscogilva 1.5–9.0 4.5 Shelley 1981
Papua New Guinea H. scabra 1.5–500 103 Shelley 1981
Papua New Guinea A. echinites 1.5–700 110 Shelley 1981
Palau were very similar. However, the catch rate for other Holothuria spp. were significantly lower. The highest average catch rates were for Actinopyga echinites
from PNG and the Solomon Islands, and for Holothuria scabra, also from PNG, with mean CPUEs in excess of 100pieces man−1hr−1. Catch rates of other species
from the Solomon Islands, Fiji, New Caledonia and PNG ranged between 10 and 40pieces man−1h−1. The catch rates for the lowest value species can be extremely
high in the earliest stage of exploitation. Adams et al. (in press) record catch rates as high as 519 pieces man−1h−1 for limited populations of H. atra in the Western
Province of the Solomon Islands, and there are even denser populations in parts of eastern Polynesia (Adams unpub. data). Of course, since these are in very shallow
water the density will quickly fall subject to fishing.
No estimates of catch or harvest rates for seaurchins were found from the Pacific islands. Mathews & Oiterong (1991) state that in Palau only a few people collect
them because of the effort required to prepare them for consumption. According to these authors several individuals working as a group can collect up to 1000 sea
urchins per day.

Two major studies on the biology and stock assessment of seacucumbers have been done in the South Pacific, in New Caledonia by Conand (1989) and PNG by
Shelley (1981). Conand (1990) and Preston (1993) have both reviewed various aspects of the biology, stock assessment, production and marketing of seacucumbers
and bêchedemer in the region. Conand (1989, 1990) describes the general biology of H. scabra, H. nobilis, H. fuscogilva, Actinopyga echinites, A. miliaris and
Thelenota ananas, including morphometrics, reproduction and growth, from the New Caledonia lagoon. Shelley (1981) conducted similar observations on
Actinopyga echinites and Holothuria scabra from around Bootless Bay on the South Papuan coast.
Both Shelley (1981) and Conand (1989) faced several problems in determining the growth
Page 476
of seacucumbers because of their plasticity, which meant that obtaining linear measures of size to obtain size frequencies was difficult, and tagging was also difficult as
tags or marks on the integument led to localized necrosis and loss within a few weeks or months. However, Shelley was able to estimate growth of Actinopyga
echinites from length size frequency data by tracing monthly modal progressions. Shelley states that a probable modal progression was also observed through part of
the data set for Holothuria scabra, but was not sufficient to fit a growth curve. Conand (1989) also estimated age and growth of Actinopyga echinites as well as A.
mauritania by tagging data, and Stichopus chloronotus and Thelenota ananas by tracing modal progressions in weight frequency distributions. Ebert (1978)
conducted a study of the growth of the small seacucumber, Holothuria atra, at Enewetak Atoll (Fig. 2, 21) by injecting individual animals with tetracycline and
measuring the increase in growth of the plates of the calcareous ring at the beginning of the pharynx. The plates of marked animals showed a flourescent mark under
ultraviolet light and the growth increment with time could be measured. This data was then used to fit a von Bertalanffy growth curve to the plate increment data, and
plate size was related to length and weight of H. atra to give a growth curve for the whole animal.
The results of these various growth estimations suggest that Stichopus chloronotus has a life span of about 5 yr, while Thelenota ananas, Actinopyga mauritania
and A. echinites have life spans in excess of 12 yr. Conand has estimated natural mortalities of populations of A. echinites, A. mauritania, Stichopus chloronotus
and Thelenota ananas in the New Caledonia lagoon and found survival rates ranged from 17 to 60% annually. Ebert (1978) estimated a natural 40% survival rate for
Holothuria atra at Enewetak Atoll in the Marshall Islands. Based on these simple data alone seacucumber populations appear capable of sustaining only light to
moderate exploitation.
Stock assessments of seacucumber populations have been made at a number of locations in the Pacific, by visual census techniques, by simply laying transects on
the reef at low water or by underwater counts using snorkel or SCUBA. The results of some of these biomass studies are shown in Table 22. Based on this limited data
set it is apparent that there are large differences in densities among locations. Some of these differences may be attributable to past and present exploitation, but as
harvest levels from different locations are unknown it is not possible to gauge the effects of removals on stock densities. That some populations can achieve very high
densities is apparent from Table 22. Ebert (1978) reports densities of H. atra at Enewetak Atoll of between 5 and 35ind.m−2, which is equivalent to between 50000
and 350000 ind.ha−1.
Some work has been done on the reproductive biology of Pacific Island holothurians, in the Solomon Islands (Holland 1994) and Guam (Richmond in press), with
the aim of developing mariculture techniques for the highervalue species, but little has yet been published.
Small subsistence fisheries for echinoderms are likely to persist in the South Pacific islands, although there may be competition from bêchedemer processors as they
turn to less valuable species with the decline in stocks of the favoured target species. As stated earlier, however, items such as seacucumbers and seaurchins are
delicacies rather than staples in the diets of Pacific islanders, so temporary declines in stock sizes and availability are not likely to have any serious effect on nutrition,
except in areas like Fiji where bêchedemer become an important food item when gardens are damaged by hurricanes. There is a considerable demand for seaurchin
roe in Asia, and Japan in particular (Conand & Sloan 1989). Special
Page 477
Table 22 Standing stock densities or biomass of commercially harvested seacucumbers in various locations in the South Pacific.
Mean species density (no ha−1)

Country Holothuria Holothuria Holothuria Holothuria Thelenota Thelenota Actinopyga Actinopyga Actinopyga Stichopus Stichopus Source
scabra fuscogilva nobilis atra ananas anax echinites Mauritania miliaris chloronotus variegatus
Papua 19.1 3.5 9.8 1.6 4.1 105.5 9.5 3 16 8.6 Lokani &
New Chapau
Guinea 1992a
(Manus)
Papua 245 54 176 211 705 31.5 1703 360 Lokani
New unpublished
Guinea
(West
New
Britain)
Fiji (Viti 2580 Preston et
Levu) al. 1988
Tonga 8.6 4.6 13.7 2.4 Preston &
(Ha’apai Lokani
Is)
Niue 182 10.7 1990
Dalzell et
al. 1993
New 683 11 13 545 18 14 847 866 512 286 Conand
Caledonia 1989
(Grande
Terre)
Page 478
ist fisheries for seaurchins have developed in countries such as Australia (Kailola et al. 1993) specifically to export seaurchin roe to Japan. There may be the
possibility of developing similar industries in the South Pacific, but careful research would be required to test consumer response to tropical species before any serious
development was considered, and the extremely high quality and freshness standards required by the Japanese market would probably make commercial development
uneconomical.
Bêchedemer fisheries in the South Pacific have been characterized by cycles of heavy exploitation followed by “fallow” periods during which the resource has had
the chance to recover. Exploitation declined in the latter half of the 19th century probably because of overfishing, whereas a slump in trade after 1930 was due to the
embargo in trade into China caused by the SinoJapanese war, followed by the continued restrictions on external trade under the Chinese communist government. The
present expansion of harvesting effort for bêchedemer in the South Pacific began in the late 1980s, partly as a result of declines in production from other bêchede
mer producers in Southeast Asia and as a result of the easing of restrictions on trade with China (Preston 1993). Bêchedemer is much sought after in China and has
apparently been useful recently as a barter item to circumvent currency transfer restrictions (Adams 1992a).
The current boom in bêchedemer export began around 1985 and was first evident in Fiji, followed by the Solomon Islands and PNG. Suddenly there appeared to
be a demand for lowervalue, easy to collect species as well as the highervalue “traditional” species (black and white teatfish, sandfish). Even though export volumes
boomed (the increase in Fiji was over 1000% in the 3 years to 1988) prices did not fall, and this has been one of the main characteristics of the bêchedemer trade:
demand is apparently insatiable or, in economic terms, highly inelastic. The volume of bêchedemer export sustained by most islands was definitely unsustainable in
the short term, and supplies are now becoming scarce, even in the remoter areas. Middlemen, often of Chinese origin, are now branching out from their bases in
Melanesia and looking at Polynesia and Micronesia as higher prices start to balance the higher cost of trade in these areas. The ease with which bêchedemer can be
harvested and the open access nature of this type of fishery means that unless there is strong intervention by government, as is the case in some locations with molluscs
such as trochus (see p. 484), then the boom and bust cycle is likely to be perpetuated.
Mollusc fisheries
Description
A large range of marine molluscs are used both for subsistence and commercial purposes in the Pacific, including bivalves, gastropods and cephalopods. Some
molluscs are harvested purely for meat, while some yield meat and shell for motherofpearl production and also pearls. Some shells, such as the cowries and olive
shells are important for the specimen shell industry, catering to tourists and conchologists (Parkinson 1984), while others in Melanesia may be burned to produce lime
for chewing with betel nut. Other uses for mollusc shells include decoration, tools such as scrapers and as traditional currency, which is still used in parts of Melanesia.
The importance of molluscs in subsistence diets in the South Pacific may be gauged by some of the shell middens that have been built up over 2000years along parts of
the South Papuan coast (Swadling 1982).
Lewis (1988b) gives a general introduction to the mollusc resources of the South Pacific and notes the importance of molluscs in the subsistence diet of some of the
islands in the
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region. Yields of 50 and 250kgha−1yr−1 are quoted by Richard (1983) for molluscs in the subsistence diets of French Polynesian atoll dwellers and those living on high
islands. Over 50 species of mollusc have been reported by Yamaguchi (1992) as being used for food and in some cases for ornamental purposes in Tonga and similar
numbers are reported to be collected from reefs and mangroves in Palau (Mathews & Oiterong 1991), PNG (Swadling 1982, Wright et al. 1983), Fiji (Rawlinson et
al. 1994) and Guam (Amesbury et al. 1986). In Fiji and Kiribati harvests of certain species constitute a significant fraction of total fisheries production. The clam
Batissa violacea is found throughout Melanesia in rivers under tidal influence, but only in Fiji is it known to be harvested in appreciable quantities with current
commercial production exceeding 1500t (in shell) annually (Anon. 1994b). Similarly, the harvests of the cockle Anadara maculosa and other bivalves in the Gilbert
Islands of Kiribati provide a cheap staple source of protein, particularly in urban areas. Bolton (1982) estimated that the total harvest of A. maculosa was over
1800tyr−1 for Tarawa Atoll, while Lewis (1988b) quotes a figure of 3,286tyr−1 for the total mollusc harvest in the Gilbert Islands.
The molluscs that are the most important foreign exchange earners for the region, exported to markets in Asia and Europe, are trochus (Trochus niloticus), green
snail (Turbo marmoratus), the blacklip pearl oyster (Pinctada margaritifera) and the goldlip pearl oyster (P. maxima). All four species yield high quality mother
ofpearl, which is used in the fashion industry to make shirt buttons and in the furniture industry for decorative inlay work. Wild pearl oysters, as their name suggests,
may also contain pearls. However, pearls can be induced to form by artificially seeding a nucleus into these oysters and this has led to successful pearl culture industries
in Australia (based on P. maxima) and French Polynesia (based on P. margaritifera). The success of the French Polynesian culture industry has led many Pacific
nations to try to develop their own pearl culture industries, but only the Cook Islands has so far succeeded with the establishment of commercial operations at Manihiki
and Penrhyn Atolls. It is notable that the northern Cook Islands is the only South Pacific island group, apart from French Polynesia, with significant remaining wild
stocks of P. margaritifera, and thus able to accumulate large quantities of natural seedstock.
The top shell Trochus niloticus was originally found in the South Pacific in Melanesia, Fiji, Wallis and the Western Caroline Islands (Palau and Yap). Because of the
demand for motherofpearl for button manufacture, trochus has been harvested extensively in the South Pacific since the start of the 20th century (Nash 1993).
Furthermore, there have been over 50 separate transplantations of trochus in the South Pacific to islands beyond its natural range in an attempt to extend the economic
benefits of trochus harvesting (Gillett 1993). Most of these transplants have generated successful fisheries in locations such as Aitutaki (Fig. 2, 1), Tahiti, Pohnpei,
Enewetak, Guam and Rota (Fig. 2, 89), and it is estimated that between 6500 and 12000t of extra trochus have been harvested over the past 50 yr as a result of these
transplantations (Adams unpub. data).
Trochus are usually found on the windward margin of coral reefs, living in the intertidal and subtidal zones (Nash 1993). The green snail or turban shell, Turbo
marmoratus, is also found in the same general environment but extends into deeper waters than trochus. Production of green snail is restricted to Melanesia,
specifically PNG, Solomon Islands and Vanuatu. Specimens from Vanuatu were successfully transplanted to French Polynesia, but unsuccessfuly to New Caledonia.
Although much less common than trochus, green snails are highly prized for their size (up to 2.0kg) and yield of high quality motherofpearl used for buttons and
especially in decorative furniture inlays. Like trochus, green snail has been commercially harvested from the Pacific islands since the start of the 20th century.
In common with bêchedemer, there is a long history of harvesting the blacklip pearl oys
Page 480
ter, Pinctada margaritifera, and the search for the elusive pearl brought many outsiders to the Pacific in the last century, particularly to eastern Polynesia (Sims 1993).
Most of the production in the South Pacific region is blacklip pearl oyster, which is most abundant in the clear shallow waters of lagoons and sheltered bays to a depth
of about 40m. The goldlip oyster is much more restricted in distribution, confined to the deeper waters of shelf areas of continents and the large islands of western
Melanesia. At present the most productive areas are the lagoons of the Tuamotu and Gambier (Fig. 2, 28) Archipelagos in French Polynesia, the northern Cook
Islands and, to a lesser extent, the Solomon Islands, Fiji and PNG. As an example of the relative productivity of these areas, the harvest of blacklip pearl shell from
French Polynesia has averaged over 450tyr−1 for the 100yr before 1980 (Intes 1986), whereas the average export of pearl shell from Fiji has been around 20tyr−1 for
the past 15yr.
Among the subsistence molluscs eaten by Pacific islanders are the giant clams (Tridacnidae), which include the largest species of all the shellbearing molluscs. As
well as being of great commercial value within and outside the Pacific region, the giant clams have attracted considerable interest because of their autotrophic nature,
relying on nutrients generated through photosynthetic symbiotic bluegreen algae in the mantle. These symbionts, known as zooxanthellae, are incorporated into the
clam mantle shortly after settlement from the larval phase. As these animals are autotrophs, it is thought that there might be considerable potential for aquaculture of
giant clams. Furthermore, in some Pacific locations, giant clam populations have been severely depleted and in some instances harvested to extinction. Reseeding from
aquaculture has been thought to offer a solution to this problem. Giant clam hatcheries have been established in Cook Islands, Federated States of Micronesia, Fiji,
Marshall Islands, Palau, Solomon Islands and Tonga (Adams & Dashwood 1994).
Most of these shellbearing molluscs are caught or harvested by simply picking them off the reef or searching in sand and mangrove mud for burrowing species. In
deeper water, fishermen can free dive for molluscs or use SCUBA gear to improve catch rates. No commercial fisheries using dredges or rakes are employed for
molluscs in the Pacific islands, although Lewis (1988b) notes that significant quantities of the scallop Amusium pleuronectes are taken in the Gulf of Papua shrimp
trawl fishery and discarded. Cephalopods such as octopus, squid and cuttlefish are also captured and used mainly for subsistence in the Pacific islands, although about
7t of octopus annually is sold through commercial outlets in Fiji (Lewis 1988b) and a similar amount in New Caledonia (Anon. 1994a). Octopus can be captured at
low tide on the reef by inserting a wire hook into their burrows and gently exciting them. Octopus, squid and cuttlefish can all be caught by spears, either from the
surface or by spear guns, and squid and cuttlefish can also be jigged on multibarbed lures used with a rod and reel.
Catch composition
Apart from Trochus niloticus, Turbo marmoratus and Pinctada margaritifera, other shells that are occasionally collected for motherofpearl are other turbo
species such as Turbo setosus and the pen shell Pteria penguin. Subsistence composition of mollusc catches has only been examined in a few locations in the Pacific
islands. Swadling (1982) reports that along the southeast coast of PNG the main mollusc species gathered for subsistence are the bivalves Geloina coaxans, Anadara
granosa, A. antiquata, Asaphis violascens, Atactodea striata, Anodonta philippina, Codakia punctata, Gafrarium tumidum, Hippopus hippopus, Tridacna
crocea, T. squamosa, Lucina corrugata, Placuna placenta and the gastropods, Strombus canarium, S. gibberulus gibbosus, S. urceus, S. luhuanus, Lambis
lambis, Littorina scabra, Turbo crassus and Nerita spp. In northern PNG Wright et al. (1983) list Strombus
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luhuanus, S. gibberulusgibbosus, S. lentiginosus, Lambis lambis, L. truncata, Nerita ornata, N. peloronta, Tridacna spp., Anadara granosa and Polymesoda
coaxans among the molluscs harvested for food in New Ireland. In the Murik Lakes region of the Sepik River delta, villagers regularly harvest the mud clams Batissa
violacea and Periglypta purpurea (Chapau 1991). GinaWhewell (1994) lists the following molluscs commonly collected for subsistence in the Solomon Islands:
Nerita spp., Chama sordida, Spondylus nicobarius, Neotia ponderosa, Tridacna crocea, T. maxima, Strombus luhuanus, Lambis lambis, Batissa fortis,
Telescopium spp., Anadara granosa, Gafrarium tumidum and Crassostrea rhizophorae.
Apart from Anadara granosa, the other regularly harvested molluscs in the Gilbert Islands of Kiribati include Gafrarium pectinatum, Asaphic violascens and
Spisula spp. (Bolton 1982). The molluscs most commonly harvested from reefs, estuaries and mangroves in Fiji include Anadara cornea, Batissa violacea,
Gafrarium tumidum, Lambis lambis, Pinctada margaritifera, Trochus niloticus, Tectus pyramis, Turbo chrysostomus and Tridacna squamosa (Rawlinson et
al. 1994). In neighbouring Tonga, the commonest species sold through commercial produce markets include Turbo setosus, T. crassus, T. argyrostomus, Lambis
lambis, Anadara antiquata, Modiolus philippinarum, Codakia tigerina, Fimbriata fimbriata, Tridacna maxima, Gafrarium tumidum and G. pectinatum. In
French Polynesia, Richard (1983) states that four gastropod species (Nerita plicata, Tectarius grandinatus, Enosania olevata, Mitra mitra) and four bivalve
species (Tridacna maxima, Area ventricosa, Chama isotoma, Cardium fragum) are the most important molluscs in the subsistence diet.
The commonest molluscs collected by women in Palau during reef gleaning are Anodontia edentula, Tridacna spp., Atactodea striata, Anadara spp., Barbitia
reeveana, Nucula rugosa, Nerita maxima and Nerita undata (Mathews & Oiterong 1991). In Guam the commonly harvested nearshore molluscs include:
Acanthopleura gemmata, Trochus niloticus, Tectus pyramis, Turbo argyrostomus, T. setosus, Nerita plicata, N. polita, Strombus gibberulus, S. luhuanus,
Lambis lambis, L. chiragra, L. truncata, Vasum turbinellus, Modiolus auriculatus, Chama spp., Ctena bella, Codakia punctata, Cardium fragum, Tridacna
maxima, T. squamosa, Quindinipagus palatum, Scutarcopagia scobotinata, Asaphis violascens, Gafrarium pectinatum and G. tumidum (Smith 1986).
According to Lewis (1988b) the cephalopods most commonly taken by coastal fishermen include the common octopus Octopus cyanea and, except in eastern
Polynesia, the lagoon squid Sepiateuthis lessoniana. The chambered nautilus, Nautilus pompilius and N. belauensis have been captured in traps set for deepslope
fishes (Blanc 1988) and in traps specifically set to catch nautilus (Saunders 1984). Despite the potential value of nautilus shells for the tourist souvenir market, this has
not led to the development of a trap fishery in any of the Pacific islands.
Catch rates
Harvesting techniques for molluscs in the coastal zone are basic and unchanged since prehistory. Emergent species lying on the lagoon floor or on coral reefs can simply
be picked up. Burrowing species in sand, silt and mud may be found with the feet and hands, or with a stick. Species attached to a rock substratum such as mangrove
oysters will need to be prised off with a knife or iron bar. Few estimates of harvesting rates for molluscs have been recorded in the Pacific from either subsistence or
commercial fisheries. Dalzell & Debao (1994) report a harvest rate of about 1.0kgh−1 for shellfish and octopus from observations on Nauru. Mathews & Oiterong
(1991) have observed catch rates in numbers of giant clams and arc shells on Palau. Catch rates of Tridacna spp. and Hippopus hippopus ranged from 9.0 to
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18.5ind.h−1, with a mean of 13.8ind.h−1, and for collection of Tridacnea crocea, 16.0 to 31.4ind.h−1 and a mean of 22.5ind.h−1. Harvest rates of mud clams in the
Murik Lakes of PNG were 1.0−2.8kgman−1day−1 or an average of 1.9kgman−1 day−1.
Lokani & Chapau (1992b) estimated catch rates for landings of trochus, green snail and black and goldlip pearl oysters over a 4yr period at Manus in northern
PNG. Their unit of effort was one fishing week, when a family unit would collect the various molluscs for cash income. Average catch rates of trochus and green snail
were 18.6kgwk−1 and 5.0kgwk−1, with catch rates of both molluscs declining over the 4yr period by between 64 and 80%. The average CPUE of blacklip oyster
remained steady at about 6.0kg wk−1 over the 4yr period while the catch rate of goldlip oyster increased from an average of 3.3kgwk−1 between 1987 and 1989 to
11.5kgwk−1 in 1990. This increase was directly related to an increase in the buying price for this oyster during 1990.
Catch rates of nautilus in Z traps in Vanuatu ranged from 0 to 2.6 nautilus trap−1 set−1, with a mean of 1.2 nautilus trap−1 set−1 (Blanc 1988), where traps were set
for between 24 and 48h. The best catch rates (2.6nautilustrap−1set−1) were experienced at depths in excess of 300m. Photographic records of trap catches of nautilus
in Palau suggest that wire mesh traps may catch in excess of 10 nautilus trap−1 (Saunders 1984). Catches were made in cubic wire mesh traps with soak times ranging
from 5 and 16h and a mean of 8h.

Few studies have been made on subsistence mollusc fisheries from the perspective of stock assessment and management. Occasional declines in Batissa violacea, or
kai, harvests in Fiji arouse concern, but, Lewis (1988b) has suggested that in general kai stocks are not being overexploited and that increased siltation in Fiji’s rivers
through agriculture and deforestation may actually be beneficial to kai populations. Adams (1992b) illustrates the possible correlations between fluctuations in kai
harvests and unusual flooding events, as well as dredging. Butler (1983) has studied some aspects of the biology and ecology of the ark shell Anadara cornea, or
kaikoso, another important subsistence mollusc in Fiji. Unlike Batissa violacea, which is essentially riverine in habitat, Anadara cornea is found in more estuarine
conditions. Butler suggested that the scale of fishing effort in parts of Fiji is such that few specimens reach maximum size due to harvesting pressure. Butler also
suggested that A. cornea populations recruit throughout the year but in a discontinuous manner and with fluctuating densities. The population biology and ecology of the
cockle A. maculosa in Kiribati has been described by Tebano (1990). Tebano considered that the absence of A. maculosa from the southern islands of the Gilbert
group may be the result of periodic droughts that affect this area causing elevated water temperatures and salinities and a decrease in dissolved oxygen beyond the
tolerance of this species. Based on studies of the reproductive biology, Tebano (1990) recommended a legal size limit for this species of 42.5mm.
Richard (1981) conducted some initial studies on the biology and abundance of four mollusc species in French Polynesia, including the smallest giant clam, Tridacna
maxima, the bivalves Area ventricosa and Cardium fragum, and the gastropod Tectarius grandinatus. Growth rates were observed from marking individuals and
measuring their growth increments whereas biomass was estimated from underwater visual census counts. The longevities of these molluscs ranged from about 3yr for
Cardium fragum to 23 yr for Area ventricosa. From the biomass densities determined from the transects, Richard (1981) determined the standing stocks of the four
mollusc species at different atolls in French Polynesia.
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The estimated density of Tridacna maxima at Takapoto Atoll (Fig. 2, 97), in the Tuamoto Archipelago, was 1400ha−1 or 67 kgha−1, while the density of the ark shell
Area ventricosa in the same location was over three times that of Tridacna maxima, with nearly 6000ha−1 or 43kgha−1 ww. In the smaller lagoon of neighbouring
Anaa Atoll (Fig. 2, 126), the density of the cockle Cardium fragrum was estimated to be 240,000ha or 460 kgha−1 ww. Similar very high densities (500000ha−1, 11
kgha−1 ww) of the gastropod Tectarius gradinatus were observed at Fangataufa atoll (Fig. 2, 24).
Because of the interest in aquaculture of giant clams, a considerable amount of study has been conducted on these species in the South Pacific during the past 15yr.
Much of this work has been reported in the proceedings of two regional workshops (Copland & Lucas 1988, Fitt 1993) and two reviews (Munro 1993, Lucas 1994)
on giant clam biology, stock assessment and aquaculture. Studies of the growth and mortality of captive and cultured clams have been conducted in a number of
locations within the region. Growth studies such as those in PNG by Munro & Gwyther (1981), in Fiji by Adams et al. (1988) and on the Great Barrier Reef (GBR) by
Pearson & Munro (1991) suggest that the longevity of giant clams is well in excess of 20yr, even for small species such as Tridacna maxima. Lucas (1994) reports
that a 500kg specimen of T. gigas was estimated to be 63 years old. Munro & Gwyther (1981) state that natural mortality of giant clams in their study (T. gigas, T.
squamosa, T. maxima, Hippopus hippopus) was extremely low and was virtually confined to small clams in the 5–15 cm range. Observed natural mortalities of
unexploited tagged clam populations on the GBR ranged from 3.4 to 10% for Tridacna gigas and 4.4% for T. derasa (Pearson & Munro 1991). Average annual total
mortalities in natural populations of T. maxima and T. squamosa in Tonga ranged from 30 to 46% and 46 to 88% respectively with means of 31 and 68% (Chesher
1991). Chesher (1991) suggests that half of the observed mortality in T. maxima was attributable to fishing, while the very high mortality of T. squamosa was due in
part to fishing and high mortalities in small (<4.0cm) specimens.
Munro (1993) states that stock densities of giant clams in the South Pacific vary enormously, attributable not only to the effects of exploitation but also to natural
causes such as episodic recruitment and juvenile habitat preference. Table 23 records some stock densities of various giant clam species in the Pacific. Stock densities
of small species such as T. maxima and T. crocea can reach very high concentrations, particularly in isolated reefs and lagoons. The figure for the extremely high stock
densities for T. maxima at Beveridge Reef (Fig. 2, 9) is the average of a range of stock densities from several different locations on the reef with a minimum of
2500ha−1 and a maximum of 200000ha−1. Beveridge Reef is an isolated oval shaped reef that is entirely awash at high tide, and encloses a lagoon of about 14km2. The
reef lies about 280km to the southeast of Niue and is rarely visited by Niueans. In some parts of Beveridge Reef the populations of T. maxima are so dense that clams
are growing on the shells of other clams. Very high concentrations of T. maxima of up to 54000 and 60000ha−1 have also been reported from parts of Aitutaki (Sims
& Howard 1988) and Takapoto Lagoon (Richard 1981), respectively, although the overall mean densities were much lower. Indeed, T. maxima in Polynesia appears
to have a markedly different community structure compared with the same species further west, where populations are much less dense, with a larger average individual
size. It is not known whether this is a genetic difference or, as appears likely, a release from competition by certain other reef borers as biodiversity decreases towards
the eastern Pacific (Adams 1988).
Descriptions of the biology of trochus are given by Bour (1990, 1992), and detailed descriptive accounts of the biology of trochus, green snail and pearl oysters in
the South Pacific and elsewhere are given by Nash, Yamaguchi and Sims respectively (all in Wright &
Page 484
Table 23 Standing stock densities of giant clams in various Pacific islands.
Giant clam densities (clams ha−1)

Location T. maxima T. gigas T. derasa T. crocea T. squamosa H. hippopus Source
Niue 89.0 14.1 Dalzell et al. 1993
Beveridge Reef 75000 Anon. 1987
Palau 1390 30–90 Hardy & Hardy 1969
Helen Reef (Fig. 2, 34) 50 33 Hardy & Hardy 1969
Cook Islands 18000 Sims & Howard 1988
Kiribati 1 1.0 1.3 Munro 1988
Great Barrier Reef 5 5 Munro 1993
French Polynesia (Takapoto) 1400 Richard 1981
Tuvalu 0.7–1.4 Braley 1988
Tokelau <6.0 Braley 1989
Vanuatu 8.9 0.14 3.5 Zann & Ayling 1988
Hill 1993). Growth of trochus has been observed from mark and recapture studies at Guam, Palau, New Caledonia and Vanuatu. Growth is relatively uniform with a
maximum life span of about 12yr and maximum sizes ranging from 11 to 15cm (shell diameter). Mortality rates have also been estimated from tagging and size
frequency data and indicate that natural mortality rates are generally low with annual survival rates of between 50 and 90% (Nash 1993), although the extremely
heterogeneous age structure of trochus subpopulations on different sections of a single reef indicate that caution must be exercised when estimating mortality rates from
size frequency data in this species (Nash et al. 1995).
Stock assessments of trochus have been conducted in a number of locations through counts along transects and in quadrats on the reef. Usually, such exercises are
carried out prior to the setting of harvest levels in islands where trochus stocks are limited such as Aitutaki and Rarotonga in the Cook Islands, and Yap and Pohnpei in
Micronesia. Trochus harvests in these locations take place over a few days or a few weeks at most and, because of the open access nature of the fisheries, a total
catch needs to be set from which quotas are allocated. In the larger Melanesian islands where the resource is not an exotic species, harvesting has traditionally
continued throughout the year with little regulation other than the setting of a minimum allowable size and, in Vanuatu and Ontong Java in the Solomon Islands,
occasional moratoria (Adams & Dalzell 1995). Nash et al. (1995) used the occasion of a trochus harvest in Aitutaki to compare three methods of estimating the
population size and hence setting a quota on the total harvest. The three methods were strip transects, the Petersen markrecapture method (Ricker 1975) and the
changeinratio (for different size classes) method (Kelber in Krebs 1989). Nash et al. (1995) concluded that the mark and recapture method was more accurate than
the strip transect for this fishery and more useful and robust than the changeinratio method. The disadvantage of this method is that the trochus population is
determined during harvesting, as tags are returned by harvesters and thus an accurate quota is not set before trochus collection begins.
Page 485
The biology and population dynamics of Pinctada margaritifera in Manihiki Lagoon (Fig. 2, 52) in the Cook Islands has been described by Sims (1992b,c).
Growth parameters suggest that like trochus, this species has a longevity in excess of 10 yr and that natural mortality rates are low. Sims (1992c) estimated an annual
natural survival rate of about 90%, with recruitment at about 9%. The relatively slow turnover of blacklip oyster populations means that they can withstand only
moderate exploitation. Sims (1992c) surveyed populations of blacklip oysters at Penrhyn and Suwarrow Atolls (Fig. 2, 95) in the Cook Islands that were overfished
in the last century and where populations have not yet recovered to former levels. Similar stock collapses have been reported from French Polynesia (Intes 1986) and
from Kiritimati Island in the Line Islands (Sims et al. 1989).
The biology of the nautilus, Nautilus belauensis and N. pompilius, has received attention. Following the discovery of a new species of nautilus in Palau (Saunders
1981), studies were done on the biology and populations of N. belauensis. Tagging studies suggested that this species grows about 3.7cm yr1 in terms of shell
increment but this rapidly decreases at the onset of sexual maturity. Saunders (1983) estimated that the average longevity of N. belauensis exceeds 20yr, and that
unlike most other cephalopods, it does not die after breeding but may live for between 5 and 10yr following maturity.
Between 1500 and 20001 of trochus were harvested annually from the Pacific islands in the period 1990–92, although this may vary considerably from year to year
depending on demand, stock sizes and the occasional moratoria that are a feature of trochus fishery management in many islands. Trochus markets are greatly
dependent on the demand for motherofpearl buttons for the fashion industry in Europe and Asia, and trends in the garment (particularly shirt) manufacturing industry
will probably dictate the future of trochus fisheries in the region. The switch from shell to plastic buttons in the 1950s caused a decline in the demand for trochus that did
not recover for two decades. It is also possible that demand (and prices), could plummet rapidly if trochus is ever promoted as an endangered species by conservation
groups. Unlikely as it sounds (since trochus is undoubtedly the bestmanaged commodity fishery in the Pacific) trochus has apparently found its way onto one
endangered species list already by being lumped together with other motherofpearl producing shells, such as green snail and goldlip pearl shell (Adams et al. 1994).
Over the past 10 years, hatchery methods for trochus aquaculture have been optimized and it is relatively easy to produce and rear juvenile specimens. It is unlikely,
however, that trochus mariculture could ever be economically feasible in the Pacific islands, but it has been hoped that cultured juveniles might be used to restock
depleted reef areas and recruitment enhancement is being tested. Experiments have been carried out in Palau, Vanuatu, Fiji and New Caledonia, but it will be some
time before it is known if these experiments were successful or not. In the meantime, the as yet untested premise that reefs can be restocked has a negative side in that
it can be cited by unscrupulous buyers and politicians as a panacea for overfishing.
Total blacklip oyster shell production from the South Pacific region currently amounts to about 400tyr1 with about twothirds of this volume being produced by
French Polynesia. Production of motherofpearl from wild stocks of blacklip oysters is confined to the Melanesian islands. Shell production from blacklip oysters in
French Polynesia and the Cook Islands is no longer from wild stocks, but from oysters cultured for pearl production. It should be stressed, however, that cultured
oyster populations originate from recruits from
Page 486
wild stocks, survival of which is enhanced through deployment on spat collectors on which the transforming larvae settle out. Closedsystem aquaculture of pearl
oysters is still in the process of being developed, but there are promising indications from a private company in Hawaii (N. Sims, Black Pearls Inc. Hawaii, pers.
comm.). The large number of sheltered lagoons in French Polynesia, coupled with naturally dense oyster populations and generous financial support from the French
Government has led to the development of an extensive pearl culture industry based on Pinctada margaritifera. By the mid 1990s over 3000 inhabitants of 40 islands
in the Tuamotu, Society, Gambier and Marquesas archipelagos derived their income from pearl culture (Anon. 1994c). The 1993 production of pearls in French
Polynesia amounted to about 2200kgyr1, worth over US$78000000, and in 1994 reputedly earned well over US$100000000 (R. Newnham, Ministry of Marine
Resources, Cook Islands, pers. comm.). Pearl cultivation is still a developing industry in the Cook Islands and at present only a small amount of pearls is produced
each year.
The success of pearl oyster cultivation in French Polynesia has generated much interest in the resourcepoor atoll microstates in the Pacific such as Tuvalu, Kiribati
and the Marshall Islands. Surveys conducted by the South Pacific Commission (Sims et al. 1989, Preston et al. 1990, Dashwood 1990) in some of the atolls of all
three states have demonstrated insufficient levels of natural stocks to support pearl culture, the result in some cases of gross overexploitation of wild stocks in the
previous and early parts of this century. Transplantation of stocks from other parts of the Pacific might be considered, although Benzie & Ballment (1994) have shown
that Pacific populations of P. margaritifera are quite highly differentiated genetically, and that care needs to taken before initiating transfers so that this genetic variation
is preserved.
Crustacean fisheries
Description
Crustaceans such as lobsters, shrimps and crabs form part of the subsistence catch in the South Pacific and form the basis of limited commercial fisheries. The trawl
fishery for penaeid shrimps in the Gulf of Papua is the only largescale commercial crustacean fishery in the region at present. Fishing techniques for crustaceans caught
for subsistence are simple and include setting traps, using nets and spears and simply collecting and gleaning from reef flats, mangroves and estuaries.
The lobster resources of the Pacific include spiny lobsters (Palinuridae) and the slipper or shovelnosed lobster (Scyllaridae). Four spiny lobster species, Panulirus
penicillatus, P. longipes, P. versicolor and P. ornatus are found on the reefs of the tropical Pacific islands. On the subtropical margins, north (Hawaii) and south
(Rapa (Fig. 2, 87), Pitcairn and Easter Island) are found P. marginatus and P. pascuensis, respectively. The slipper lobster resources in the South Pacific are less
well documented but Parribaccus caledonensis, P. antarcticus, P. holthuisi and other Parribaccus spp. are found on the tropical reefs of the Pacific in the same
general locations as the spiny lobsters. On the subtropical margins are found species such as Scyllarides haani and S. squammosus which, like the spiny lobsters, are
commercially valuable.
A feature of the behaviour of the four tropical spiny lobsters is their apparent reluctance to enter traps or pots, which are used to great effect with subtropical and
temperate spiny lobsters elsewhere in the Pacific and beyond. Prescott (1988) reports that traditionally manufactured traps have been used to capture Panulirus
penicillatus and P. longipes in various
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parts of the Pacific, using bait such as chitons and seaurchins. The introduction of underwater torches and dive gear has turned Pacific island fishermen away from
traditional methods of catching spiny lobsters. Furthermore, commercially manufactured fish traps, used to catch P. marginatus in Hawaii and P. cygnus in Western
Australia have been tried in many parts of the Pacific with little success. Consequently, the basis of all commercial and subsistence fishing for lobsters in the Pacific
remains diving with spears and handnets, and collecting on reef flats at night during low tides. Innovations such as the use of SCUBA or hookah gear may be
introduced to improve the efficiencies of dive fishermen but essentially the methods remain similar throughout the region. Certain practices, such as the use of tangle nets
on the forereef, and the use of compressed air to flush lobsters out of holes, are generally discouraged as they may be considered destructive of reefs.
Zann (1985) gives a good description of a smallscale commercial lobster fishery in Tonga that targets P. penicillatus. Fishermen are active in the week before and
after new moon, or between 15–18 nights per month. Divers work at night in pairs, starting at about 20.00h, usually working until 04.00h. The divers systematically
work a given area of reef then unload their catch, change torch batteries and continue searching until 04.00h when fishing is terminated and the catch is landed at a
processing facility. Grandin & Chauvet (1994) and Chauvet & Farman (1994) describe lobster fisheries of Lifou (Fig. 2, 49) and Isle de Pins (Fig. 2, 36) in the French
territory of New Caledonia. Originally, P. penicillatus was caught at Lifou using traps made from plant fibres baited with seaurchins, but as is the case elsewhere, trap
fishing has been superseded by nighttime dive and spear fishing. This is also the case in the Cook Islands where lobsters (mainly P. penicillatus) are captured by reef
walking at night either side of low tide or free diving on reefs with torches at night (Passfield 1988).
At Yule Island (Fig. 2, 119) in the Gulf of Papua, fishermen dive during the day to catch P. ornatus by hand (Prescott 1988). Coral bommies are also surrounded
by tangle nets to snare lobsters when they are frightened from their refuges. Gill nets set to catch fish at Yule Island will also take P. ornatus. The same species is the
target of Australian and PNG fishermen in the Torres Straits. Divers work singly or in pairs during the day catching P. ornatus by hand or with short spears (Kailola et
al. 1993). An unusual feature of the P. ornatus fishery of the Torres Strait and Gulf of Papua is that this species is susceptible to trawls. P. ornatus undertakes a long
spawning migration from reefs in the Torres Strait at the western edge of the Gulf to Yule Island in the east. Crossing the Torres Strait and the Gulf of Papua they walk
through areas fished by commercial shrimp trawlers. For several years in PNG, this lobster bycatch was seen as a useful seasonal bonus by the trawl fleet, but
eventually was banned owing to concerns about the reduction of spawning stocks and the concomitant effects on subsequent recruitment of P. ornatus to the northern
GBR and Torres Strait.
The portunid crab Scylla serrata is found throughout most of the tropical Pacific wherever there are areas of muddy substratum with stands of mangrove trees
(Brown 1993). Like spiny lobster it has been caught by Pacific islanders for subsistence purposes and forms the basis of small commercial fisheries in some countries of
the region. Mangrove crabs are caught in the South Pacific in baited wire mesh traps or, more commonly, by carefully pulling crabs from their burrows with a metal or
wooden hook. If crabs can be extracted carefully without damage to the burrow then the vacated burrow will soon be reoccupied and crabs can continue to be
captured at the same site for many years (Swamy 1994).
In Fiji, mudcrabs are caught using both methods. Along the drier north and western zones of Viti Levu, crabs are caught in the mangroves mainly from burrows. A
piece of mangrove
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stick is inserted into the burrow and is seized by the crab. The stick and crab are carefully removed from the burrow that is blocked on extraction of the crab to prevent
escape and reentry. According to Swamy (1994), the fishers who regularly catch crabs in this fashion know the locations of several burrows within a mangrove
system. These locations are well guarded secrets as they will continue to produce crabs if not damaged during harvesting activities. On the wetter south side of Viti
Levu, crabs are caught predominantly in dilly traps set in the intertidal mangroves. The dillies are made of a gill net fastened to a circular steel hoop. They are baited
with meat or fish and set in mangrove streams at low tide. Unlike pots or cage traps, the dillies are not left unattended, but raised regularly to see if there are crabs
feeding on the bait.
Some pot fishing is also conducted in Fiji, where wire basket traps are left baited and unattended, and emptied every 6h, although according to Swamy (1994), this
method is not very popular among local fishermen. This method of crab fishing is used in Palau (Nichols 1991) as well as more traditional methods such as hand
collection and spearing. Capture of mudcrabs on Pohnpei is mainly by fishers searching for them in the swamps, looking for crabs in burrows, hiding in between
mangrove roots or submerged in puddles (Perrine 1978). Some crabs are also caught on sandy reefs up to a kilometre from the mangroves, hiding in seagrass and
seaweeds. Crabs are also caught this way in Guam but the predominant method of capture is with baited dillies and in traps (Amesbury et al. 1986). In New Caledonia
a range of methods are used to catch mudcrabs including traps, removal from burrows and, occasionally, seine nets (Delathiere 1990).
Penaeid shrimp capture fisheries are not of any economic significance except in PNG where the Gulf of Papua trawl fishery produces between 1000 and 1300t of
shrimps annually. Fishing is conducted in waters between 5 and 40m deep with little fishing on the deeper slopes. Shrimp fisheries elsewhere in PNG and the Pacific
islands are more modest. The Orangerie Bay fishery on the southeast Papuan coast generates between 5 and 781 annually (Anon. 1994d). Along the north coast of
PNG penaeid shrimps have been captured by smallscale beam trawls in the Markham (Quinn & Kojis 1986) and Sepik estuaries (Frusher 1985a, Chapau 1991).
Similar exploratory fishing in the seagrass beds adjacent to the Nggurambusu River (Fig. 2, 68) (Guadalcanal) in the Solomon Islands revealed populations of Penaeus
semisulcatus, including significant numbers of gravid females (Delaune 1989). Although it was thought not economically viable to fish this stock commercially, it might
form a valuable source of brood stock for shrimp farming and culture.
Smallscale shrimp fishing is conducted in Fiji, with an annual production of between 3t and 5t (Choy 1988), although there are occasionally years with higher
catches on unusually dense spawning aggregations (M.Lagibalavu, Fisheries Division, Fiji, pers. comm.). Shrimps are caught with spears, hand nets, scissor nets and
seines (Choy 1981, 1988). Gill net fishing in the lagoon of Tongatapu, the main island of Tonga, generates a small bycatch of penaeid shrimps, mainly P. semisulcatus
and Metapenaeus ensis. Braley (1979) conducted a series of beam trawls in Tongatapu lagoon to assess the feasibility of this smallscale method of catching shrimps
and to collect information on the principal species in the subsistence catch.
Other crustaceans that are collected mainly for subsistence purposes in the Pacific include the three spot reef crab (Carpilius maculatus) and the red reef crab
(Etisus splendidus). Amesbury et al. (1986) state that these crabs are captured by hand on reefs in Guam. Dalzell et al. (1992) also reported subsistence consumption
of these crabs on Niue. Land crabs are also important throughout the Pacific, particularly the common purple land crab Cardisoma spp. These species swarm at
certain times of the year in a spawning migration to release eggs
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in the sea and can simply be collected in large numbers by parties driving along roads at night and picking them up. The coconut or robber crab, Birgus latro, may be
very common in some Pacific islands and is caught in substantial quantities in Vanuatu, both for subsistence, but more recently for the local tourist industry (see
contributions in Brown & Fielder 1991).
The burrowing mud lobster, Thalassina anomala, is commonly found in mangrove areas of the Pacific and is caught in appreciable quantities in Fiji, where it is
regarded as a delicacy. The lobster is caught in a snare trap inserted into the burrow and triggered by the movement of the lobster. Burrowing mantis shrimps
(Stomatopoda) that live in seagrass meadows are also caught for subsistence by snare traps. The snare is inserted into the burrow attached to a baited stick that
protrudes from the burrow. When the shrimp attacks the bait the stick vibrates and the snare is tightened by the fisher. An even simpler method for mantis shrimp
capture has been reported from New Britain in PNG (M.Chapau, Dept. of Fisheries & Marine Resources, PNG, pers. comm.) where bait is loaded into a nylon
stocking that is pushed into the burrow. The shrimp attacks the bait but gets its claws tangled in the stocking and can then be extracted from the burrow.
Some mention should also be made of the freshwater prawns, Macrobrachium spp. Although normally found in fresh water, M. rosenbergii uses the mangroves of
the Gulf of Papua as a nursery and berried females may be caught in waters with salinities as high as 15–18‰ (Frusher 1983). Furthermore, many of the Pacific islands
rivers are short, and freshwater species such as Macrobrachium are caught in the vicinity of the coastal margin. A description of traditional fisheries for freshwater
prawns on the Sepik River of PNG is given by Robertson (1983). Macrobrachium can be caught on baited hookandlines or in traps and the methods described by
Robertson for PNG are similar to those used elsewhere in the South Pacific. Although only a minor item in the diet of most Pacific islanders, Macrobrachium prawns
have attracted interest for their aquaculture potential.
As with finfish, the crustacean fauna of the outer reef slope has attracted some interest owing to the presence of pandalid shrimps and geryonid crabs, both of which
are catchable in traps. King (1993b) summarized the information on pandalid shrimps and surveys of populations in the Pacific. It was initially hoped that in resource
poor countries without extensive estuarine areas, trap fisheries based on pandalid shrimps might develop as another source of income for fishermen. This hope has only
been realized in Hawaii, however, and these fisheries have not persisted because of poor returns from catches (Polovina 1993). Trial fishing with traps in both Palau
(Hastie & Saunders 1992) and the Marquesas Islands in French Polynesia (Poupin et al. 1991) have revealed populations of the geryonid crabs, Chaceon granulatus
and Chaceon species nova that are readily caught in crab pots. Both surveys suggest that populations might support smallscale fisheries.
Catch composition
Apart from the lobster fisheries of Yule Island and the Torres Strait, the dominant lobster in catches in the South Pacific islands is Panulirus penicillatus (Pitcher
1993). The data presented in Table 24 from various locations in the Pacific confirm the dominance of P. penicillatus throughout much of the region. The next most
important species would appear to be P. longipes. Exact numbers of species at different locations were not always obtainable, for example Pitcher (1993) states that
P. ornatus makes up virtually 100% of the Torres Straits lobster catch, although P. versicolor is not uncommon. Similarly, the Yule Island lobster catch is composed
almost entirely of P. ornatus with the occasional capture of
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Table 24 Percent species composition of spiny lobster catches from various locations in the South Pacific region.
Location P. penicillatus P. longipes P. versicolor P. ornatus Source

Lifou (New Caedonia) 80.0 15.0 4 1 Grandin & Chauvet 1994
Isle de Pins (New Caledonia) 60.7 38.8 0.4 0.1 Chauvet & Farman 1994
Niue 72.5 25.4 2.1 Dalzell et al. 1993
Yule Island (PNG) <1.0 >99.0 Anon. 1984b
Torres Strait (PNG/Australia) <1.0 >99.0 Pitcher 1993
Tsoi Islands (PNG) 98.0 1.0 1.0 Hair & Aini 1994
Kiribati 90.7 9.3 Prescott 1977
American Samoa 100.0 Saucerman 1994
Western Samoa >99.0 <1.0 King & Bell 1991
Fiji 95.5 1.4 3.1 Richards 1994
Tonga 67.9 27.5 4.6 Prescott 1993
other species. King & Bell (1991) examined over 1000 spiny lobsters in Western Samoa between September and November 1989 and found only two specimens of
P. versicolor, with the remainder being P. penicillatus. Saucerman (1994) lists only P. penicillatus from American Samoa and this appears to be the only species
present.
According to George (1974) and Pitcher (1993) the distribution of spiny lobster species is a function of several physical variations, such as coral cover, wave
energy, water quality and turbidity. P. penicillatus is largely restricted to the reef crest and reef slope, preferring oceanic water that is clear and well oxygenated by
constant wave action, and habitat with plenty of coral cover for protection. The same general habitat is preferred by P. longipes, but this species can be found in
deeper waters and extends on to the reef plateau into more sheltered waters. P. versicolor ranges from the reef crest and slope to the well developed coral gardens on
the reef plateau. P. ornatus is found in the widest range of habitats from sheltered, moderately turbid waters of backreef lagoons to the very turbid silty areas near
rivers and mangroves. It is the commonest species on broad continental shelves and can be found at a depth >200m.
Between 50 and 60% of the shrimp catch in the Gulf of Papua shrimp fishery consists of a single species, Penaeus merguiensis, or banana shrimp. Other common
catch components are tiger shrimps (P. monodon, P. semisulcatus), which form about 12% of the catch, and endeavour shrimps (Metapenaeus spp.) constituting a
further 20% (Gwyther 1980). Banana shrimps dominate the Orangerie Bay shrimp fishery in southeast Papua, where they form between 80 and 90% of landings, with
the balance of the catch including Penaeus monodon, P. semisulcatus and Metapenaeus spp. (Anon. 1994d). About half the landings of experimental catches of
shrimps in Milne Bay was Penaeus merguiensis, with a further 22% comprising P. monodon and P. semisulcatus (Coates et al. 1984). Commercialscale shrimp
trawling surveys around the mouth of the Sepik River were dominated by banana shrimps (P. merguiessis, P. indicus), which formed over half the catch with another
40% comprising endeavour shrimps (Metapenoeus affinis, M. ensis, M. dobsoni, M. demani) and the balance made up from tiger shrimps (Penaeus monodon,
P.japonicus, P. semisulcatus) (Frusher 1985a).
Shrimp catches by smallscale beam trawling in the Markham River estuary were almost entirely Metapenaeus demani (Quinn & Kojis 1986), with occasional
amounts of Penaeus
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semisulcatus. Shrimp catches using a similar type of gear along the coast of Guadalcanal caught only the giant black tiger shrimp, P. monodon (Delaune 1989). Small
scale otter trawling in the Tongatapu lagoon caught almost equal quantities of the tiger shrimp, P. semisulcatus (46%) and the endeavour shrimp, Metapenaeus ensis
(54%) (Braley 1976, 1979). In Fiji, the dominant shrimp captured by smallscale fishermen using spears and nets is Penaeus canaliculatus, which forms 85% of the
catch around Suva (Choy 1988). The balance comprises a mixture of P. latisulcatus, P. monodon, P. semisulcatus, Metapenaeus anchistus and M. elegans.
Catch rates
Catch rates for spiny lobsters are infrequently reported from most of the Pacific and are not expressed in a uniform manner. Catches may be recorded as numbers or
weight, and may be expressed per hour, per day or per trip. For example, the average catch rate for spiny lobsters reported by Grandin & Chauvet (1994) from the
New Caledonian island of Lifou is 3.2kgh−1, but 12.1kgday−1 was reported from the Isle de Pins (Chauvet & Farman 1994). Even where catch statistics are reliable,
problems occur because improvements in technology have increased fishing efficiency. Catch rates of dive fishermen based at Daru catching lobsters in the Torres Strait
prior to 1985 ranged from 6.0 to 13.9kg−1man−1day−1, with a mean of 7.9kg−1 man−1 day−1. The introduction of hookah gear has led to a major increase in CPUE
that peaked in 1989 at 36.3kg−1 man−1 day−1 (Anon. 1994d). Catch rates in the Tongan fishery during the early and mid 1980s ranged from 0.5 to 4.9kgman−1h−1
(mean= 1.5kgman−1h−1) or 2.4 to 16.4 kg night−1 (mean=8.5 kg night−1) (Zann 1985). Lobster catch rates from several atolls in the Gilbert Islands ranged from 0.6
to 21.3lobsterman−1h−1 or a mean of 11.3lobsterman−1h−1 (Prescott 1977). Assuming a mean weight similar to lobsters in the neighbouring Marshall Islands (Ebert &
Ford 1986) of 0.45kg, then these catch rates translate to 0.27–9.6kgman−1h−1, or a mean of 5.1kgman−1h−1.
Unlike the lobster fisheries of the tropical Pacific, catches of the Hawaiian lobster, Panulirus marginatus, in the commercial lobster fishery of the subtropical
Northwest Hawaiian Islands were made by trapping in plastic lobster pots. Average annual catch rates in the Northwest Hawaiian Islands commercial fishery,
expressed as lobsters trap−1 haul−1, declined from 2.75lobsterstrap−1haul−1 in 1983 to 0.56lobsterstrap−1 haul−1 in 1991, with an overall mean of
1.4lobsterstrap−1haul−1, or 1.65kgtrap−1haul−1 to 0.34kgtrap−1haul−1 with a mean of 0.84kgtrap−1haul−1 (Polovina 1993). About 80% of the catch is formed by P.
marginatus, with most of the balance comprising the slipper lobster, Scyllarides squammosus. A small recreational dive fishery for Panulirus marginatus at Midway
Atoll (Fig. 2, 59) was monitored between 1975 and 1978 by MacDonald (1987). Catch rates were expressed as lobsters diver−1 trip−1 and varied from 0.6 to
4.5lobstersdiver−1trip−1 with a mean of 2.8 lobsters diver−1 trip−1, or 0.53 to 4.0kg diver−1 trip−1 with a mean of 2.5 kg diver−1 trip−1, based on the mean lengths of
male and female lobsters in the catch and lengthweight equations given by Uchida et al. (1980).
Clarke & Yoshimoto (1990) report the results of an intensive depletion fishing experiment on an unexploited stock of Scyllarides squammosus on Laysan Bank
(Fig. 2, 48) where catch rates for legal sized lobsters ranged from 1.2 to 3.61obsterstrap−1haul−1 and a mean of 2.341obsterstrap−1haul−1. A small percentage of
slipper lobster catches in the Northwest Hawaiian Islands is formed by the slipper lobster S. haani. This species comprised the majority of lobsters caught in traps
around the Pitcairn Islands group in the extreme southeast of the South Pacific region. Average catch rates ranged from 0 to 8.1 lobsters haul−1 with
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a mean of 1.54 lobsters trap−1, or 0 to 9.3kg and a mean of 1.35kg from fishing at Oeno, Ducie and Henderson Islands (Dalzell 1994, Sharpies 1994). The best
fishing conditions were experienced at Ducie Island, where catch rates ranged from 0.7 to 8.1lobstershaul−1 (mean=2.75 lobsters haul−1) or 0.81 to 9.3kghaul−1
(mean=3.14kghaul−1). The maximum and minimum catch rates mark respectively the initial and final CPUEs from the start and finish of a 15 dayperiod of intensive
fishing. This decline, however, is more likely to be the influence of the lunar periodicity, with catches declining markedly at around the time of the full moon towards the
end of the fishing period. Other species taken in small quantities included Panulirus penicillatus and P. pascuensis
As with the spiny lobsters, catch and catch rate data for mudcrabs in the South Pacific are sparse and not uniform. Swamy (1994) describes methods of catching
mud crabs in Fiji but does not include information on CPUE from traps or by harvesting crabs from burrows. Commercial catch statistics for crab catches from Palau
are relatively crude and refer to catch per trip. Nichols (1991) reports that between 1984 and 1990 the CPUE ranged from 8 to 22kgtrip−1 with a mean of 14kgtrip−1
or roughly equivalent to 14 crabstrip−1. Mathews & Oiterong (1991) report an average catch rate of 0.7crabstrap−1 or 0.7kgtrap−1 from nightly trapping for mudcrabs
in Palau during new moon by a single individual. Perrine (1978) gives no information on catch or harvest rates from the traditional mudcrab fishery on Pohnpei but does
include some information on trials with wire and plastic crab traps set in the mangroves. The average catch rate from these experiments in two separate trials was about
0.6 kg trap−1. Delathiere (1990) reported an average catch rate from New Caledonia of 5.08 kg ind−1 day−1 for commercial harvests during the permitted fishing
season (February to November), mainly from women fishers catching crabs directly from their burrows.
Mudcrabs are caught by hand in the Murik Lakes, northern PNG, with the CPUE for commercial harvesting ranging from 1.4 to 6.2kgind−1day−1 or a mean of
3.6kgind−1day−1 (Chapau 1991). Chapau (1991) suggested that subsistence catch rates were likely to be lower, in common with those in the Purari Delta at the apex
of the Gulf of Papua, which ranged from 0.0 to 0.08kg ind−1 day−1. (Haines 1978/79). Trial catch rates in the same area with chickenwire crab pots ranged from 0.2
to 1.25 crabs trap−1 or 0.08 to 0.5 kg trap−1 for a 4h soak (Anon. 1980). Gill nets and beach seines were also tested, and returned catch rates of 2 to 3 crabs per 1
hr soak and 0.4 to 0.75crabs per haul respectively, or 0.8 to 1.2kgsoak−1 and 0.16 to 0.3kg haul−1 (Anon 1980). The most effective method of capture, however,
was by hand and stick and averaged about 5.5 crabs man−1 h−1 or 2.2kgman−1h−1. Matsuoka & Kan (1989) reported on the catches made with different trap types
set in mangrove swamps adjacent to the Markham river mouth. The most productive design was a hemispherical trap, with an entrance at the top. The trap caught an
average of 0.4crabstrap−1 or in terms of weight, 0.14kg trap−1.
Catch rates of all shrimp species in the Gulf of Papua trawl fishery ranged from 11.5 to 24.1kgh−1 with a mean of 15.8kgh−1 between the years 1974 and 1993
(Evans & Opnai 1994a). Catch rates for the banana shrimps were 6.0–15.2kgh−1 (mean=8.3kgh−1) during the same period. Catch rates for all shrimps in the
Orangerie Bay fishery in southeast Papua showed a much greater range (8.0 to 35.8kgh−1) and a higher mean CPUE (27.0kgh−1) between 1981 and 1993. More
modest shrimp catch rates were encountered during the trawl survey of Milne Bay (Coates et al. 1984), where the CPUE range was 1.2 to 11.6kgh−1 with a mean of
6.2kgh−1. Three commercial shrimp surveys in the Sepik River mouth encountered catch rates ranging between 0.0–15.6kgh−1 with an overall mean of 6.4kgh−1
(Chapau 1991), similar to those for Milne Bay.
Catch rates for smallscale beam trawling in the Markham River mouth are not given by
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Quinn & Kojis (1986), but a mean CPUE of 0.7kgh−1 can be computed for Metapenaeus demani, which made up most of the shrimp catch. Catch rates from the
deployment of similar gear in and around the Murik Lakes in the Sepik Delta produced catch rates ranging from 0.0 to 1.89kgh−1 or a mean of 0.5kgh−1 (Chapau
1991). Braley (1976, 1979) does not give catch rates in terms of kgh−1, using instead the numbers caught per standard 20min period of trawling. However, the data
are sufficient to reestimate the CPUE in terms of weight, and based on his data this ranges from 0.3 to 2.4kgh−1 with a mean of 1.0kg h−1. Choy (1988) reports
average monthly catch rates from combined spear and handnet fishing in Laucala Bay (Fig. 2, 46), Fiji, in terms of numbers per minute for juveniles and adults. Taking
combined catches and raising these to catch per hour gives a range of 78 to 222 shrimps h−1 or a mean of 124shrimpsh−1. Assuming an average shrimp weight of 2.3g
(Choy 1981) then this translates to an average CPUE of about 0.29kgh−1.

Considerable attention has been given to the biology and stock assessment of the two lobster species that constitute the Northwest Hawaiian lobster fishery, Panulirus
marginatus and Scyllarides squammosus. The biology of both species has been summarized in Uchida & Uchiyama (1986). The catch and fishing effort data for this
fishery, based on log books kept by commercial fishermen, were used to estimate the MSY and optimum fishing effort using a dynamic production model (Schnute
1977). The model provided a good description of the fishery between 1983 and 1989, but then catch rates declined markedly in 1990 and 1991. Polovina (1993)
stated that fishing effort alone was not sufficient to cause these declines and that instead the decline is the result of poor recruitment attributable to oceanographic
conditions (Polovina et al. 1994) at some banks, which resulted in a concentration of fishing effort at the remaining banks where recruitment was strong. More recent
research has revealed that recruitment of Panulirus marginatus at two of the more productive banks in the Northwest Hawaiian Islands is related to relative sea level
height, which is in turn linked to the ENSO event (Polovina & Mitchum 1992).
Less attention has been given to the slipper lobster, Scyllarides squammosus, which constitutes an average of between 10 and 50 per cent of lobster landings in the
Hawaiian fishery. As stated earlier, Clarke & Yoshimoto (1990) conducted an intensive fishing experiment to obtain biomass estimates through localized depletion and
application of the Leslie method. The biomass estimate and catchability coefficient obtained from the intensive fishing experiment were then used to estimate the
biomass from other banks and islands where fishing had been conducted and there were data on catch rates.
A detailed descriptive summary of the biology of Panulirus penicillatus is given by Prescott (1988) and Pitcher (1993). P. penicillatus populations have been
studied in Palau (MacDonald 1982, 1988), the Solomon Islands (Prescott 1988), Marshall Islands (Ebert & Ford 1986), Tonga (Zann 1985, Prescott 1990, Munro
1992), Western Samoa (King & Bell 1991) and New Caledonia (Chauvet & Farman 1994, Grandin & Chauvet 1994). Growth of this species has been determined
from tagging data (Enewetak, Solomon Islands) and from length frequency data (Tonga, Western Samoa, New Caledonia), suggesting life spans in excess of lOyr. The
abundance of P. penicillatus has been determined in the Solomon Islands and Enewetak by mark and recapture experiments and from Lifou (New Caledonia) by
cohort analysis (Grandin & Chauvet 1994). Relative densities of P. penicillatus in the Solomon Islands ranged from 111 to 128lobsterkm−1 of reef edge
(mean=120lobsterskm−1) and between 35 and 164 lobsters km−1 (mean= 126 lobsters km−1) in Enewetak. Grandin & Chauvet
Page 494
estimated a total fishable stock of P. penicillatus at Lifou of 410000 lobsters. A rough estimate of the coastline length of this island and hence reef edge is 207km,
which suggests a relative density of 1980 lobsters km−1, or nearly 20 times those observed in the Solomon Islands and Enewetak.
Ebert & Ford (1986) used yield per recruit analysis to estimate the optimum harvest size for P. penicillatus at Enewetak, suggesting that 40% of the adult
population could be harvested annually. Adams & Dalzell (1993) have used the relative biomass estimates of Ebert & Ford (1986) and Prescott (1988) to suggest that
an annual harvest rate of 20kg of lobster km−1 of reef may be sustainable and should be used as an initial approximation or rule of thumb when trying to determine the
size of lobster resources on a given island or reef. The disparity between biomass estimates in Lifou and Enewetak and the Solomon Islands may be due to differences
in methodology or indeed reflect the much higher productivity of the P. penicillatus population at Lifou. Grandin & Chauvet (1994) note that the annual harvest of
lobsters from Lifou is about 15000 lobsters, which represents only 3.7% of the fishable stock. These few data serve to show the uncertainty that still exists over the
scale of Pacific spiny lobster resources.
Growth rates of P. longipes from Tonga (Munro 1992) and New Caledonia (Chauvet & Farman 1994) have also been determined from length frequency data and
suggest maximum life spans of over 5yr. Some aspects of the biology of P. versicolor from Palau, mainly reproductive biology and fecundity (see Pitcher 1993), have
been given by MacDonald (1982, 1988).
A considerable research effort has been conducted on the populations of P. ornatus in the Torres Strait and the Gulf of Papua. During the 1970s and 1980s, it was
established through tagging programmes that the lobsters that formed the basis of a seasonal fishery at the eastern edge of the Gulf at Yule Island between November
and March originated from reefs in Torres Strait (Moore & MacFarlane 1984, Bell et al. 1987). Indirect evidence from the seasonal appearance of lobsters in the Gulf
of Papua trawl fishery indicated the Torres Strait as the likely point of origin. MacFarlane & Moore (1986) suggested that the migratory population that spawned on
arrival at Yule Island is the major source of larval recruitment in the western Coral Sea, including the east coast of Queensland and Torres Strait. These findings led to
the imposition of a ban on trawling for lobsters in the Gulf of Papua. Other aspects of the biology of this species from the Torres Strait, such as growth and mortality,
are given by Pitcher (1993).
A synopsis of the biology of Scylla serrata and a review of the limited amount of study conducted within the region is given by Brown (1993). Mudcrab biology and
populations within the South Pacific have been studied in Pohnpei, Fiji, New Caledonia and PNG. Swamy (1994) gives an account of two populations of mudcrabs on
the southeast and the northwest coasts of Fiji. The two locations are physically distinct in that the northern site experiences about half of the rainfall (1500mm) normally
falling in the south. Growth, mortality and recruitment parameters for both populations were described from length frequency data. The biology of S. serrata in New
Caledonia has been examined in detail by Delathiere (1990) to provide information for managing this resource. Estimates of the total habitat occupied by mudcrabs
were determined using remote sensing techniques, and the biological observations, including growth, mortality, recruitment and feeding, were made on captive and wild
populations. Aspects of the biology of S. serrata in Pohnpei and in Guam are described by Dickinson (1977).
The biology and stock assessment of mudcrabs in PNG have been described for populations in the Gulf of Papua (Frusher 1983, Opnai 1986b) and in the Murik
Lakes (Chapau 1991). Densities of crabs at two sites in the Purari Delta ranged from 10.4 to 21.4crabsha−1 or 2.5 to 4.7kgha−1, with means of 16crabsha−1 and
3.6kgha−1 respectively. In the Murik Lakes,
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mudcrab densities in four sites ranged from 13 to 39crabstrap−1 with a mean of 23 crabs trap−1 or 3.6 and 8.5 kg ha−1 and a mean of 6. 0kg ha−1. Perrine (1978)
estimated that a minimum estimate of the crab standing stock on Pohnpei was 3000 crabs in a mangrove area of 5400 ha−1 or 0.6 crabs trap−1 or 0.25kgha−1. Perrine
suggested that the true density may be two to three times this estimate. Delathiere (1990) estimated the abundance of mudcrabs in mangroves in New Caledonia to be
7900crabstrap−1 or a biomass of 1.04tha−1. As with the spiny lobsters, there appears to be major differences between the biomass estimates of crabs in New
Caledonia and the other Pacific islands. This may be due to much greater densities of mudcrabs in New Caledonia or, as is more likely, differences in sampling
techniques and the size and locations of sample areas. Delathiere (1990) counted crabs in a relatively small area (48m2) most of which were juveniles. Density estimates
in Pohnpei and PNG were based on counts of adults made over much larger sampling areas of several hectares.
The biology and stock assessment of shrimps in the Gulf of Papua are addressed in studies by Gwyther (1980, 1982), Tenakanai (1980), Branford (1982), Frusher
(1985b), Frusher et al. (1985), Polovina & Opnai (1989), Waffy (1990) and Evans & Opnai (1994a,b). The biology and ecology of the most important species in the
fishery, the banana shrimp Penaeus merguiensis, has been described by Gwyther (1980), Frusher (1983, 1985b) and Frusher et al. (1985). Aspects of the biology
and ecology of the endeavour shrimps (Metapenaeus spp.) are described by Tenakanai (1980), Frusher (1983) and Waffy (1990). Gwyther (1980, 1982) made the
first serious attempts to estimate the MSY from the Gulf of Papua shrimp fishery using the Fox (1970) surplus production curve to model catches for the whole shrimp
fishery and for catches of Penaeus merguiensis only. Gwyther’s analysis considered only 6yr of data (1974–1979) to estimate an annual MSY for the shrimp fishery
of 1200t and 570t for catches of banana shrimps only. Evans & Opnai (1994a) fitted a Fox curve to catch and effort data for the years 1974 to 1993 to determine a
total fishery MSY of 1500t and a banana shrimp MSY of 620t. Evans & Opnai suggested that the maximum economic yield for the fishery is between 1350 and 1400t
with an optimum fleet size of 15 vessels.
Polovina & Opnai (1989) showed an inverse relationship between the minimum monthly rainfall between September and December and Gulf of Papua shrimp
catches in the subsequent year. When a monthly rainfall term was incorporated into the Fox production curve, more of the variation in catch could be explained than by
an effortonly production model. Polovina & Opnai (1989) suggested that heavy rainfall may create extensive areas of low salinity water extending beyond the coastal
mangroves that cause the postlarval shrimps to settle out before reaching their nursery grounds within the mangrove environment. Alternatively, heavy rainfall may
create a constant outflow from the mangroves, negating any inshore tidal flow and preventing postlarvae from being carried inshore. The inverse relationship with catch
and rainfall is the opposite of that observed in the banana shrimp fishery in the neighbouring Gulf of Carpentaria in northern Australia (Staples 1985), where the
correlation between rainfall and subsequent fishery catches is positive. However annual rainfall in the Gulf of Carpentaria is about 900mm compared with 2700mm in
the Gulf of Papua. Evans & Opnai (1994b) have made a detailed examination of the relationship between shrimp CPUE and rainfall. They suggest that the true
relationship between rainfall and yield of banana shrimps is parabolic rather than linear, with an optimum annual rainfall of around 1200mm. They suggest that low
rainfall is critical immediately following the major annual postlarval settlement, and during the period of offshore recruitment in February to March.
The biology of the witch shrimp Penaeus canaliculatus in Laucala Bay, Fiji has been described by Choy (1981, 1988). Choy used length frequency data to
estimate growth and mortality and described the reproductive biology and recruitment of this species. The stand
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ing stock of P. canaliculatus was determined from Beverton & Holt’s (1957) swept area method and this was used in turn with the natural mortality rate to compute
potential yield. Some observations on the biology of P. semisulcatus and Metapenaeus ensis from Tongatapu Lagoon are reported by Braley (1979). Catches of the
two species were inversely correlated but the CPUE for both species was reduced during the full moon. Both species undertook spawning migrations outside the
lagoon between January and March.
The biology of deep water pandalid shrimps in the South Pacific region has been summarized by King (1987, 1993b), and includes details of those countries where
resource surveys of deep water shrimps have been conducted. Stock assessment of pandalid shrimp populations have been made in the Northern Mariana Islands
(Ralston 1986) and Hawaiian Islands (Ralston & Tagami 1992) using Leslie depletion fishing methodology. Estimates of standing stocks ranged from 2.4kgha−1 in
Hawaii to 5.5kgha−1 in the Northern Mariana Islands. Both studies suggest that there was a limited resource of pandalid shrimps and that this could be easily over
exploited by trap fishing.
The biology and stock assessment literature on other crustacean resources in the South Pacific includes the coconut crab from Vanuatu, where it is economically
important in the tourist trade (see contributions in Brown & Fielder 1991). A synopsis of the biology of this species in the South Pacific region is given by Fletcher
(1993). No other biological or stock assessment studies appear in the literature on other land crabs in the South Pacific region. Some aspects of the biology of the deep
water geryonid crabs from Palau and the Marquesas Islands are given by Hastie & Saunders (1992) and Poupin et al. (1991).
The lobster fisheries of the South Pacific continue to require divers to capture animals because of poor performance using pot or traps. The dominant species
comprising most of the catch in Pacific island lobster fisheries, other than the southern Papuan coast, will undoubtedly be Panulirus penicillatus, which occupies a
relatively narrow band of the outer reef crest. This habitat area is limited in many Pacific island countries because of the small size of most of the islands in the region.
Despite these obvious limitations, a number of ventures have tried to establish commercial lobster fisheries on a relatively large scale in a number of Pacific countries,
usually without much success. Adams & Dalzell (1993) discussed the apparent and true potential of lobster fisheries based mainly on P. penicillatus. They catalogued
a history of failed commercial enterprises in the region that had not taken account of the true economic potential for these fisheries.
Some smallscale operators do derive incomes from P. penicillatus fisheries, such as in northern PNG where buyers collect lobsters from village fishermen and air
freight them to restaurants and hotels in urban areas (Hair & Aini 1994). Overheads are kept to a minimum and these smallscale fisheries are able to return a profit.
The P. ornatus fishery in the Torres Strait is also based on divers spearing lobsters, but fishermen are able to sell into the more lucrative Australian market where
prices for fresh lobsters range from 30 to 70 A$kg−1, far more than could be realized in most of the Pacific islands. The introduction of hookah gear into this fishery
markedly increased catch rates and doubled the volume of the fishery from 30–40tyr−1 to 70–80tyr−1. The effects of such an obvious increase in efficiency in the P.
ornatus fishery will only be known in the long term. Commercial fishing interests have approached Pacific island governments with the intention of using hookah gear to
harvest P. penicillatus. Given what is thought to be the limits of the resource, this increase in efficiency would probably only lead to more rapid depletion of limited
stocks.
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Fisheries for shrimps outside of PNG are clearly very limited, but there is growing interest in mariculture of shrimp in the South Pacific region. Mariculture of penaeid
shrimps has been successfully developed in New Caledonia, with annual production of over 600t. The cultured stock is not from the South Pacific but is Penaeus
stylirostris, originally imported from the USA (Eldredge 1994). A smaller amount of maricultured shrimp (≈60tyr−1) is produced in French Polynesia, where
Macrobrachium spp. are also cultured with an annual production of between 15 to 20t (Anon. 1994c). Fiji has several smallscale penaeid shrimp farms with a total
annual production of around lOt. These were based, until recently, on postlarvae imported from Australia but there are now increasing quantities of postlarvae coming
from local hatcheries (Anon. 1994b).
As with spiny lobster fisheries, mangrove crab fisheries will probably continue to be smallscale enterprises, given that most production in the South Pacific region is
probably from collection rather than trapping. Live lobsters and crabs command much better prices and there may be potential for some small export fisheries to parts
of Asia to cater for specialized highquality, highvalue markets for live seafood. This has already been demonstrated from the Melanesian islands for selected reef fish
species, and trial airshipments of live spiny lobster have been sent from PNG to Hong Kong (Richards 1993). However, local prices are often sufficiently high that
export is not an economically attractive option, particularly in areas where overseas tourists are common.
Shrimps, mudcrabs, coconut crabs and spiny lobsters are the most commercially valuable crustaceans in the South Pacific. Other edible resources such as mud
lobsters and land crabs have some local market value but are not in demand worldwide. However, large immigrant Pacific island communities are found in New
Zealand, Hawaii, Guam and Australia, and can generate considerable demand for traditional delicacies, including items such as fish and shellfish. For example, Schiller
(1989) estimated that the annual export of coconut crabs from Niue in 8 months between 1987 and 1988 amounted to between 3200 and 5900 animals. All these
crabs left Niue as passenger luggage en route to New Zealand, where the majority of Niueans now live. There is enormous demand for coconut crab in Asia also, and
stocks of this species in every part of the Pacific can now be considered depauperate, if not endangered. Other resources such as pandalid shrimps have attracted initial
government enthusiasm for their development, but the resource has failed to meet expectations and the only commercial fisheries, in Hawaii, have ceased production.
Similarly, it is likely that fisheries on deep water geryonid crabs are not economically feasible in the region and, if they do become so in future, Pacific island
governments would be wise to have management measures in place for what promise to be fisheries with a low sustainable exploitation rate (Adams & Dalzell in press).
Coastal fisheries production

Having reviewed the various sources of coastal fisheries production we attempt here to estimate the volume of landings from various coastal fisheries and to estimate the
value of this production. As far as possible, estimates of landings have been taken from annual reports, technical reports or other papers published between 1989 and
1994. Information on landings prior to 1989 was used in some instances where data from recent landings were not available. Most countries have some estimate of
commercial fisheries production and these are usually published in some form of technical document. Furthermore, estimates of total fisheries pro
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duction for most countries are included in the FAO Yearbook of Fisheries Statistics. These figures range in quality and accuracy and may include landings from offshore
tuna fisheries. Accurate estimates of subsistence fisheries production were usually not available. In some cases subsistence fisheries production had to be estimated
from nutritional data as the product of per capita food consumption and population size. In the few instances where no estimate of national subsistence fisheries
production was available at all (PNG, Tuvalu, Western Samoa), then the FAO figure was used (FAO 1994) as a nominal total, and subsistence production estimated
as the difference when known commercial landings were subtracted. The various sources used for each country are listed in Table 25. Many of the sources used here
fall in to the “grey literature” category, i.e. departmental technical memos, unpublished manuscripts and in some cases simply personal communications from individuals
who have access to production records. The annual fisheries production summaries for each Pacific island country
Table 25 Sources of data used to estimate the volume and value of national subsistence and commercial coastal fisheries production for the South Pacific
Islands.
Country Data sources and notes

American Samoa Commercial and subsistence production from Hamm et al. 1994 and Craig et al. 1993.
Cook Islands Subsistence catch based on nutritional data in MategnaSmith et al. in press and J. Dashwood (SPC Fish. Prog. pers. comm.). Commercial finfish
production in 1988 from Zoutendyk & Paulo 1989. Trochus landings from Nash et al. 1995 and pearl shell production from Cook Is unpublished
records
Federated States of Subsistence catch in based on nutritional information in Elymore et al. 1989. Commercial landings adapted from Smith 1992a
Micronesia
Fiji Subsistence and commercial production based on Anon. 1994b
French Polynesia Subsistence and commercial production from Anon. 1994c
Guam Commercial and subsistence production from Hamm et al. 1992, 1994
Kiribati Commercial and subsistence production from Anon. 1991c
Marshall Islands Commercial catch figures from Smith 1992b, subsistence production based on nutritional data in Anon. 1991e
Nauru Subsistence and commercial production from Dalzell & Debao 1994
New Caledonia Subsistence and commercial production from Anon. 1994a
Niue Subsistence and commercial catches based on Dalzell et al 1993
Northern Marianas Commercial fisheries production in Hamm et al. 1994, Subsistence fisheries prodution based on nutritional data for urban Micronesians in Elymore et
al. 1989
Palau Commercial and subsistence production from Anon. 1993b
Papua New Guinea Total coastal fisheries production from FAO 1994, commercial landings from Anon. 1989
Pitcairn Island Subsistence fisheries production based on average monthly numbers of fish reported between 1973 and 1992 in Pitcairn Miscellany, the island’s
newsletter
Solomon Islands Subsistence and commercial production figures from Skewes 1990 and Richards et al. 1994. Note about 2,500 tyr−1 of bait fish (mainly anchovies
and sprats) also caught in coastal lagoons by domestic poleandline fleet Nichols & Rawlinson 1990
Tokelau Subsistence fisheries production based on observations by Hooper 1983
Tonga Total coastal fisheries production from Anon. 1991f, commercial finfish production in Munro 1990 and commercial lobster production in Zann 1985
and Prescott 1990
Tuvalu Total coastal fisheries production from FAO 1994. Commercial fisheries production from Patiale & Dalzell 1990 and unpublished
Vanuatu Total coastal fisheries production from David & Cillauren 1989, commercial fisheries production in Anon. 1991g, Anon. 1992b and Anon. 1992d
Wallis & Futuna Total coastal fisheries production from Anon. 1994e and commercial fisheries production from A.Ledreau pers. comm.
Western Samoa Total coastal fisheries production from FAO 1994 and Zann et al. 1991, commercial fisheries production on Upolu from Helm 1992
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Table 26 Mean annual coastal fisheries production in the South Pacific 1989–94.
Catch Weight (t) Value (US $)

Commercial reef and deepslope fish 10414 27,258964
Commercial coastal pelagic species 4252 14,028423
Commercial estuarine fish 1688 3,417745
Commercial crustaceans 1839 14,250593
Commercial bêchedemer (processed to 10% fresh wt) 1604 10,070966
Other echinoderms 30 31087
Commercial trochus, green snail, pearlshell 2495 10,995145
Other molluscs 2003 1,747741
Total commercial catch 24325 81,800664
Total subsistence catch 83914 179,914623
Total coastal fisheries catch 108239 261,715287
from the sources listed in Table 25 are given in Appendix 1 (p. 510). These data sources were used to compile the annual catch volumes and values for subsistence and
commercial landings in each of the countries and territories of the region and these are included in Table 25.
A summary of the totals in each sector is given in Table 26 and the subsistence and commercial catches by country are summarized in Table 27. For some countries
such as Fiji, New Caledonia and French Polynesia, the annual catch refers to the most recent annual report of fisheries statistics, usually 1993 or 1994. For most other
countries the production figures are a mix of information on various resources from various years between 1989 and 1994 as noted in Table 25. The total coastal
fisheries production in the South Pacific during
Table 27 Mean annual subsistence and commercial production from coastal fisheries for the countries and territories of the South Pacific between 1989
and 1994.
Country Subsistence fisheries Nominal Value Commercial fisheries Value Total fisheries Nominal Value
production (t) (US$) production (t) (US$) production (t) (US$)
American Samoa 215 814238 52 178762 267 993000
Cook Islands 858 3,047683 124 314761 982 3,362444
Federated States of 6243 11,237400 637 1,483544 6880 12,720944
Micronesia
Fiji 16600 45,767395 6653 18,340043 23253 64,107438
French Polynesia 3691 14,468720 2352 14,371469 6043 28,840189
Guam 472 1,935632 118 456413 591 2,392045
Kiribati 9084 13,373667 3240 4,770000 12324 18,143667
Marshall Islands 2000 3,103213 369 714504 2369 3,817717
Nauru 98 219600 279 628605 376 848205
New Caledonia 2500 9,000000 981 3,968650 3481 12,968650
Niue 103 471504 12 54720 115 526224
Northern Marianas 2825 12,280427 141 613804 2966 12,894231
Palau 750 1,805192 736 2,410059 1485 4,215251
Papua New Guinea 20588 41,176000 4966 22,096 908 25554 63,272908
Pitcairn Islands 8 16000 0 0 8 16000
Solomon Islands 10000 8,405660 1,150 4,343811 11150 12,749471
Tokelau 191 104509 0 0 191 104509
Tonga 933 1,901208 1429 2,806641 2362 4,707849
Tuvalu 807 657781 120 97811 927 755592
Vanuatu 2045 1,953360 467 1,514364 2512 3,467724
Wallis & Futuna 621 3,105360 296 2,316729 917 5,422089
Western Samoa 3281 5,070074 208 319066 3489 5,389140
Total 83914 179,914623 24327 81,800664 108241 261,715287
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the early 1990s is estimated to be about 108000tyr−1, worth US$ 261715287 (Table 26). The population of the South Pacific region is about 6500000 people giving
an annual per capita coastal production of 16.6kg. If the large inland population of 2700000 in PNG is discounted then the annual percapita production is about
28.4kg. This coastal fisheries production estimate is not equivalent to fish consumption, since it does not take account of canned fish imports, tuna consumption, or
exports. Depending on the island group, actual seafood consumption will tend to be at least 25% higher than domestic coastal fishery production, because of the status
of canned mackerel as a nonperishable rural staple, and much higher than this in highly developed, denselypopulated islands like Guam that have limited domestic
catches.
In the South Pacific 80% of the catch from inshore fisheries, whether from reefs, estuaries or fresh water, is estimated to be taken for subsistence purposes with the
remainder (20%) being directed to commercial markets. Most of the commercial finfish goes to domestic markets, and most of the commercial invertebrates are
exported. Subsistence fisheries production was estimated to be about 83900tyr−1, and would have been worth US$179900000 if sold in domestic markets. In most
instances, the value of the subsistence catch is based on the average figure for commercial fish landings. Commercial fisheries landings amount to about 24300tyr−1,
worth US$81800000. Not included in this total is the production from aquaculture which, among other items, would add another US$78000000 from the production
2200kg of pearls from French Polynesia in 1993 and US$7280000 from shrimp farming in New Caledonia during the same year.
Discussion
In this contribution we have tried to document the characteristics of fisheries in the coastal zone of the South Pacific islands, on reefs, deep reef slopes, the pelagic zone,
and estuaries. We have made some mention of freshwater production which is significant in both Papua New Guinea and Fiji, by the volume of finfish caught and,
especially in Fiji, by the volume of molluscs harvested. In both locations most of the freshwater finfish catch comprises exotic species such as tilapia and common carp
(Cyprinus carpio). With respect to the coastal fisheries resources, our focus has been largely on the fisheries rather than on the biology and ecology of the target
species. However, relevant biological literature for work accomplished in the Pacific is also reviewed. Many of these resources are common to neighbouring Southeast
Asia and as far to the west as East Africa. A few extend across the Pacific to the western coast of Central America, while others have a circumtropical distribution.
However, we have generally omitted references to studies outside of the South Pacific in order to document fully the work conducted on coastal species in the region.
Information on the comparative biology of coastal fish and invertebrate species within and outside the region is given in many of the reviews contained in Wright & Hill
(1993) and the various literature cited therein. Similarly, the reviews of reef fish biology in Sale (1991), Polovina & Ralston (1987) and Polunin & Roberts (1996) take
a global perspective and contain comparative information on fish populations and species on AtlantoCaribbean, Indian Ocean and Pacific reefs and reef slopes.
Most of the countries of the South Pacific are by definition developing economies, which implies low industrial output, a small proportion of the population involved
in manufacturing and, in some countries, drift to urban centres of landless poor who are obliged to fish for income and sustenance. The amount of fisheries research
conducted in most locations is
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minimal and the administrations of most South Pacific countries are not well versed in modern methods of fishery management (Adams & Dalzell in press). It will be
clear from this paper that, with the possible exception of PNG, the focus of most fisheries research in the region is in developed countries, or in overseas territories
where economies are supported by metropolitan administrations. Thus much of the work quoted here has been conducted in northern Australia, New Caledonia,
Hawaii, French Polynesia and Guam. All these are locations where there has been considerable effort expended on marine biological research, especially on coral reef
species, although much of it is not directly applicable to fisheries stock assessment and management. Unfortunately, it is the poorer states in the region, with few
economic opportunities and populations increasing at between 2–3% per year, that tend to develop unsustainable fisheries, and that urgently require improved
quantitative information about the status of their coastal fisheries (Adams et al. 1995).
The present state of knowledge about most of the coastal fisheries resources represents a fraction of what is known about the same or similar resources in
neighbouring South and Southeast Asia. However, the accumulation of information on the biology of coastal fisheries resources in the South Pacific has accelerated,
particularly over the past 20yr. The factors responsible for this accumulation include a greater awareness of the importance of coastal fisheries resources, improved
recruitment and training of national fisheries biologists in some countries, introduction of new techniques for studying tropical fish populations and the realization that
some old established techniques used in temperate water fisheries are also applicable in the tropics. The advances made during the past 20yr in electronic micro
processor technology have made computers readily available to fisheries biologists throughout the region. Apart from the convenience of being able to manage large
databases on a daily basis, a whole range of different fisheries stock assessment applications has been developed by several institutions. Although not designed
exclusively for tropical fisheries, the motivation to develop these programs has often arisen from experience in trying to analyze data from tropical fisheries.
Conventional approaches to estimating population parameters through age and growth data developed for temperate fisheries often have not been applicable in the
tropics.
This problem led initially to development of lengthbased stock assessment methodologies that were in turn incorporated into many of the computer applications. The
basis for several of these routines was the estimation of growth parameters from length frequency data and subsequent estimation of other life history information such
as mortality rates, recruitment and relative or absolute yields. Such algorithms are based on the premise that growth of a cohort or year class can be traced through
successive length frequencies, usually monthly samples, and that growth curves can be fitted to the length data through some iterative routine given seeded values of the
curve parameters. Many of these length basedmethods are included in Pauly & Morgan (1987) and are reviewed by Rosenberg & Beddington (1988). Useful though
lengthbased methods are, they have not received universal acclaim and have frequently attracted much justified criticism when applied without taking their constraints
into account (see for example Hilborn & Walters 1992). One major problem has been the inability of several of these programs to calculate confidence limits around
the estimates that they generate. However, lengthbased methods have proved effective with shortlived species, particularly the small pelagic fishes used for tuna bait.
These specialized fisheries software packages allied to commercial software developments in database, spreadsheet and graphical programs have made a significant
contribution to the expansion of information on coastal stocks in the South Pacific.
Another important development in tropical fisheries science that had an influence on fish stock assessment in the South Pacific region was the appreciation that the
microstructural
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increments observed in fish otoliths were, in most cases, laid down on a daily basis and so provided a means of ageing fish when annuli were not discernable (Campana
& Nielson 1985). Various examples are given in the previous sections of fishes aged through otolith microstructure. In all but the smallest species, the otoliths must be
sectioned or ground to the required thickness and polished to display the growth surfaces by transmitted light, and further treatment such as acid etching and vacuum
coating with gold are required for scanning electron microscopy. Where the fish have reached a considerable age and the otoliths are dense and crystalline, the record
of growth is incomplete leading to the method proposed by Ralston (1981) where incomplete counts are used to interpolate mathematically the entire growth record.
The various studies of ageing reef, deep slope and small pelagic fishes using otolith microstructure have been documented above.
In temperate water species the distinct seasons during the year are reflected in the growth record of fishes in the form of yearly annuli on scales, bones and the
otoliths. Ageing of fishes is therefore relatively easy and fisheries administrations in Europe and the USA routinely age many tens of thousands of fish each year to
determine the age structure of the population. Ageing fish through observations of the yearly increments on the otoliths and scales has not been seriously attempted in
many locations in the region, usually due to the assumption that tropical species do not possess yearly annular marks. However, an increasing body of evidence is
emerging to suggest that yearly annuli are present in otoliths and scales of tropical fishes and these have been validated from New Caledonia (Loubens 1980, Coutures
1994), the GBR (Walker 1975, McPherson 1992, McPherson & Squire 1992) and from Hawaii (MoralesNin & Ralston 1990). While these are all on the
subtropical margins of the South Pacific, there is also evidence that yearly annuli are detectable in fishes such as nemipterid breams (Chapau 1993), emperors (Dalzell
et al. 1992) and snappers (Dalzell unpubl. data) in nearequatorial locations. Current research in northern PNG is in the process of validating the periodicity of annular
marks in some of these species (Chapau pers. comm.).
Hillborn & Walters (1992) state that age estimation is the most important information obtained from sampling fish catches and in this review we have narrowed our
focus to those studies concerned mostly with age and allied parameter estimation such as mortality rates. Other aspects of life history, such as reproductive biology,
food and feeding behaviour, have not received much attention and information on these for most of the species and resources discussed here can be found in Polovina
& Ralston (1987), Sale (1991) and Wright & Hill (1993). If ageing of many of the coastal fishes in the Pacific is achievable through the use of long established simple
techniques for reading yearly annuli then this will be a major step forward for coastal finfish stock assessment in the region. However, one other major area of ignorance
in the fisheries biology of most of the coastal fisheries in the South Pacific (and indeed in most of the other oceans) is the relationship between stock and recruitment.
This is fundamental when deciding the allowable exploitation rate for a stock of marine organisms.
This question is particularly pertinent to the commercial invertebrate fisheries of the region, especially giant clam, green snail, pearl oyster and coconut crab, where
capture is straightforward and populations can be depleted to the point of extinction. As stated earlier, there are a number of examples of pearl oyster populations in the
South Pacific that were severely depleted between 50 and 100 years ago and have not recovered. The present boom in bêchedemer processing means that
holothurian populations are being reduced markedly in inshore waters, with little understanding of the recruitment processes or recovery rates, although most bêchede
mer species and trochus are probably proof against ultimate extinction given their cosmopolitan nature and cryptic habits. Other possible examples of recruitment over
fishing include the Torres StraitYule Island lobster population and deepslope fish
Page 503
stocks on seamounts. The failure of the annual spawning migration to Yule Island following trawling on previous migrating stocks led to a ban on catching lobsters in
trawls in 1984. The ban has been maintained since then and the Yule Island fishery has recovered. The highly concentrated populations of deepslope snappers and
groupers on seamounts can be fished down very rapidly without the possibility of recovery from new recruitment. The Tongan deepslope fishery rotates between
different seamounts, leaving areas that have been heavily exploited to recover (Latu & Tulua 1991) while seamounts that have been left unexploited have accumulated
fishable populations. The process of recruitment of snappers and groupers to seamounts is poorly understood. Some seamounts when depleted do not recover within a
few years such as the Haputo Pinnacle at Guam, where deepslope stocks were fished to virtual extinction in the late 1960s (Ikehara et al. 1970) and have never
recovered.
Investigation of stock and recruitment for coastal fish populations in the South Pacific islands has been confined to small pelagic species, namely Encrasicholina
heteroloba in Palau (Muller 1976) and E. heteroloba, E. devisi and Spratelloides gracilis in PNG (Dalzell 1984a). Both authors attempted to fit two basic stock
and recruitment models developed by Ricker (1954) and Beverton & Holt (1957) to data generated by bait fisheries in these two locations. Both authors found that
although these simple models would explain some of the variation in recruitment with respect to parental stock biomass, other factors such as rainfall and standing stock
biomass of other species also influenced recruitment (Muller 1976, Dalzell 1984a,b). Recruitment of coral reef species has attracted a great deal of interest, mainly from
ecologists, and several relatively longterm investigations have been conducted on reef fish recruitment on the GBR. Much of this work has been conducted on small
economically unimportant species, such as damselfishes and small wrasses, that can be easily observed underwater. Recruitment to a particular reef with respect to
spawning stock biomass may be confounded by the planktonic larval stages of most reef species, which means that they may be dispersed far from their natal reef
(Doherty 1991). This has led to one school of thought that believes that biomass densities of fish on coral reefs are mainly a function of recruitment and that post
recruitment processes are not of any great importance (Doherty 1991). Conversely, other workers have acknowledged the impact of recruitment on reef fish
populations but also regard postrecruitment processes as equally important in determining abundance (Jones 1993). From a management perspective it is unlikely that
for Pacific island multispecies reef fisheries recruitment processes will ever be sufficiently understood to be incorporated into detailed management initiatives.
In giant clams, it is likely that there is no smooth relationship between stock and recruitment, but rather an abrupt cutoff below a certain parental density. Giant
clams maximize fertilization success by spawning in synchrony in response to currentborne pheromones, thus there is likely to be a certain population density below
which synchronous spawning becomes unlikely and fertilization becomes rare. However, even in dense natural populations of giant clams producing, presumably,
enormous quantities of fertilized eggs, recruitment is very sporadic and appears to depend far more on environmental conditions and available settlement sites than on
parental stock biomass (Adams et al. 1988, Pearson & Munro 1992).
One area where a great deal more work needs to be done is on the “catchment areas” for recruitment of different species and communities. Some species, such as
trochus and giant clams, have shortlived larvae and are probably locally recruited, at the level of the individual reef. The restricted natural distribution of trochus,
despite its proven ability to become established on almost every reef to which it has been introduced, is an indication that natural longdistance transport of trochus
larvae is not common. Yet the larvae of other species may be carried in the plankton from country to country and even from region to region
Page 504
before settlement. Arising from observations on the rate of recovery of damaged reefs, there are indications that certain Western Australian corals and reef fish
populations may be dependent on Indonesian sources for replenishment (D.Williams, Australian Institute of Marine Science, pers. comm.). The parameters affecting
recruitment will obviously have a large bearing on the approach to managing different fisheries—there may be little point in trying to maintain fishable trochus stocks
through the medium of one large reserve, but rather through a series of small reserves on the local scale. There may be little point in maintaining a reserve at all if it
cannot be placed upstream of a target (“larval sink”) area. For coastal species with longerlived larvae it may be necessary for management to be coordinated on a
national, or even regional, basis if it is to be effective.
It will be clear from the data presented in this review that catch rates of artisanal fishermen in the coastal zones of the South Pacific islands are very low, usually of
the order of a few kilogrammes for several manhours of work. Only the trawl fishery in the Gulf of Papua catches fish in amounts comparable to other tropical shelf
fisheries in Asia but retains only the shrimp element of the catch, dumping the greater finfish component back into the sea. Catches of bait fish by the other industrial
scale fishery in the region, the poleandline tuna fishery, are modest, amounting to between 100 and 200kg night−1. However, even with limited smallscale fishing
gears, people can have a severe impact on fish and invertebrate populations, where numbers of fishers are sufficient to generate high fishing mortalities. Russ (1991) has
noted that even when populations are not driven to extinction, the effects of fishing pressure may modify the behaviour as a response to heavy fishing pressure in
shallow reef areas. Russ cites as an example parrotfish, normally associated with the lagoon and reef flat in the Philippines, moving away from their preferred habitat
down the reef slope into deeper waters to escape fishing pressure. In the Pacific, Johannes (1981) states that following the widespread spear fishing at night of
Bolbometopon muricatum in Palau, this species sought dormitories away from the shallow reef flat on the deep reef slope. Dalzell & Debao (1994) report that
fishermen on Nauru believe that spear fishing has had a similar effect on shallow reef populations of snappers and groupers, which have retreated into deeper waters
through constant fishing by SCUBA equipped spear fishermen. Furthermore, Nauruan fishermen report that spear fishing has markedly reduced snapper and grouper
populations and that large reef snappers and groupers are seldom seen. The decline in catches from reefs in the lagoon of Woleai Atoll following the widespread
introduction of underwater spear fishing prompted a ban on this fishing method (Smith & Dalzell 1993). Spear fishing is not conducted with SCUBA gear at Woleai but
fishermen equipped only with goggles can be highly efficient at reducing fish populations on reefs.
New types of fishing gear may be introduced to an island and rapidly become very popular, so much so that fishing effort rapidly increases but without sufficient
documentation to assess the effects on reef and lagoon fish populations except at a gross level. Yeeting & Wright (1989) review such a development in Tarawa Atoll,
Kiribati, during the 1970s and 1980s, but were only able to make fairly crude analyses based on collective catch and effort. Fishermen in Kiribati have become
concerned about the decline in bonefish (Albula glossodonta) catches. This has occurred over the period of the expansion in gill net fishing but is also confounded by
the construction of causeways between the atoll islands that have had a deleterious effect on the spawning migrations of this species. Whether this decline in stocks is
due chiefly to overfishing or to environmental effects through causeway construction is unknown, although recent data on the bonefish stocks certainly suggest that
survival of this species in Tarawa Lagoon is at a critical stage with no recently observed spawning migrations (Anon. 1993d). Management under these circumstances is
particularly difficult,
Page 505
given that bonefish is a traditional subsistence food on Tarawa, and there is little information to judge whether fishing or causeway construction has the greatest negative
influence on bonefish populations.
The population of the South Pacific presently amounts to around 6.5 million people, 62% of those living in PNG. Of the 4 million people in PNG, only onequarter of
the population lives on the coast. The total coastal population of the region is therefore approximately 3.25 million people, equivalent to the population of a medium
sized city in Europe or the USA, but spread over an area of 29 million km2 of ocean. Most of the countries and territories of the South Pacific are thus heavily
dependent on aid to maintain their economies. The typical island state is small and it has few natural resources. Trade among islands is also limited because what one
island can offer, namely the products of subsistence farming and fishing, can be found on most other islands. Islands that produce cocoa, palm oil and copra are
competing against much larger producers elsewhere in the tropics. Some manufacturing enterprises do exist but are small and generally serve limited domestic markets.
Furthermore, imports of raw material tend to be costly because relatively small quantities have to be shipped over long distances.
Probably the greatest economic potential for many of the South Pacific islands, particularly those with limited natural resources, is tourism (Anon. 1991d). In some
countries such as Fiji, Vanuatu, and parts of Micronesia, tourism is a major source of revenue and employment. Other countries in the region are trying to develop a
better tourist industry infrastructure and increase earnings from this sector of the economy. The growth in the tourist trade may also offer greater potential for coastal
fishermen as this will provide an additional market for their products and possibly a chance to demand a higher price than may be realized by domestic markets.
However, the negative aspects of this growth are that quality fish and invertebrates may be channelled in increasing volumes away from domestic markets, and demand
for specialist products may lead to largescale stock depletion. A good example is Vanuatu, which receives about 45000 tourist visitors per year. Among the
attractions offered to visitors are gourmet dining on local produce such as flyingfox, deepslope fishes and coconut crabs. The coconut crab, in particular, is one of the
main items on the menus of restaurants catering for tourists. The downturn in other incomeearning opportunities in rural areas, such as copra, has led to an increase in
the harvesting of coconut crabs in Vanuatu, to the point where concern about localized extinctions led to an Australian funded research project to gather management
information (Brown & Fielder 1991).
The other major influences on coastal fisheries resources and future fisheries development in the South Pacific are likely to come from neighbouring Southeast and
Northeast Asia. Already, the majority of motherofpearl is sold to markets in Japan and Korea, while the Chinese have established a virtual monopoly on the buying of
bêchedemer. While the coastal fisheries of Japan are highly regulated and managed, this is not entirely true of the countries of Southeast Asia, where combinations of
high population growth and in some cases poverty, have created large artisanal fisheries that continue to cater for a growing demand for fish (Pauly 1989). Part of this
demand is for highquality reef fish and, as discussed previously, this has generated the development of live reef fish fisheries in a number of countries in the west of the
South Pacific. Many of the countries of the region still have traditional systems of marine tenure and regulation of coastal fisheries, although this tends to break down
where population pressure is very high and urbanization increases. However, Crocombe (1994) suggests that one of the major challenges to fisheries management and
marine tenure will be the increasing influence of Northeast and Southeast Asia. If the pressure from Asia to share in the exploitation of the region’s coastal resources
increases, this is
Page 506
bound to cause conflict with traditional tenure systems and may create serious political problems for the countries of the region.
Fisheries development initiatives in the South Pacific over the past 50 years have sought to improve fishermen’s incomes by improving catch rates, identifying more
lucrative markets and increasing the value of the landed catch through teaching better postharvest techniques. Unfortunately, very few development initiatives for an
exportoriented capture fisheries in the coastal zone have had any success. One typical example of this process was the establishment of fisheries on deep reef slope
stocks, where success was very shortlived in most areas where it was tried. Handline catch rates of snappers and groupers of the deep slope are on average two to
three times greater than the equivalent effort spent in shallow reef fishing (Table 5, p. 418), and might be as high as ten times on virgin stocks. As the deepslope
fisheries developed, it was clear that domestic markets alone would not support these fisheries. This led to the search for more lucrative markets overseas in Japan,
Hawaii, Australia and New Zealand. Hawaii in particular was the marketing focus of the two biggest deepslope fisheries in the region, Fiji and Tonga, and the opening
of this market led in turn to better handling and preservation of fish to realize the best prices. Because this was an auction market, with a limited capacity, Fiji and Tonga
exporters found themselves having to compete against each other.
The negative side of the development of deepslope fisheries in the South Pacific was that they demonstrated most of the mistakes commonly associated with
fisheries development in the region. The catch rates encountered on fishing virgin stocks were often used by fishermen to compute economic returns from fishing for
deepslope species although catch rates at MSY are likely to be onethird to onehalf of those at the start. The products of deepslope fishing, large snappers and
groupers, are not the most favoured fish of Pacific islanders, who prefer the smaller more pungently flavoured shallow reef fishes to the more delicately flavoured
species from the deep reef slope. The development of the deepslope fisheries in locations like Tonga and American Samoa was also driven in] by easy credit and soft
loans to fishermen for purchasing fishing dories (Itano 1991, Latu & Tulua 1991), with the intention of stimulating expansion of the fishery before the true limits of
productivity were known. These fisheries then became overcapitalized, with excessive numbers of vessels competing for a shrinking resource, with the inevitable result
that fishermen were forced to default on loan repayments and many vessels turned to other activities (such as, in Fiji, harvesting seacucumbers) or simply ceased
operations. Similarities with patterns of fisheries development elsewhere are strong (Smith 1994).
As discussed earlier, the deepslope fisheries have declined in favour of mediumscale fisheries for the larger tuna species caught and exported, usually fresh, to
Japan and Hawaii for the lucrative sashimi trade. However, the experiences of developing the overseas markets for unfrozen deepslope fishes were valuable when
establishing similar markets for unfrozen large pelagic species. Commercial finfishery development initiatives in the region, particularly export initiatives, are likely to
continue to turn towards oceanic pelagic stocks rather than coastal fisheries because of the much larger unexploited resource base. The successful fisheries in Fiji, and
elsewhere in the region, are proving the viability of adapting monofilament longline gear to small and medium sized, locally based, fishing vessels and the highquality
fish produced by these South Pacific fisheries appears to be maintaining its market share in Japan and the USA in the face of competition from the large number of small
Chinese longliners now operating in the north of Micronesia. The main commercial limiting factor is transporting the catch by air from the Pacific islands to overseas
markets. Some countries such as Palau, French Polynesia, New Caledonia, Fiji, Federated States of Micro
Page 507
nesia and Tonga have direct air links with Japan and Hawaii. Other countries where pelagic fisheries are developing will have to try to find some way of accessing these
overseas markets and take advantage of the better returns on landings.
For the smaller resourcepoor countries of the Pacific, bêchedemer and motherofpearl shell fisheries offer only reducing potential for generating income, given the
lack of sustainability as a management objective in most areas. Shallowwater invertebrate fisheries are easily overexploited and any income is generally considered to
be “windfall” cash for village families rather than a basis for permanent investment. These fisheries are also strongly influenced by external market forces beyond the
control of national economies and could easily go into price decline through decreased demand, as happened with trochus in the late 1950s, and again in 1991.
Management of sessile invertebrate harvests, particularly in smaller countries, would have to be severely limited to maintain sustainable yields. Present management
strategies for trochus in the former US Trust Territories of Micronesia and the Cook Islands are based on very short harvest seasons with quotas dictated by pre
season stock assessments. If a large part of the population expects to share in the revenues from resources such as trochus then limited seasons with short pulses of
intensive effort may be the only way to effectively manage these stocks. It has also been suggested for bêchedemer (R.Richmond, University of Guam, pers. comm.)
that producers form a cartel and rotate harvests between islands each year, giving depleted stocks the chance to recover in those islands not participating in an annual
harvest, or even that trochus and bêchedemer harvest periods be opened alternately to provide a similar recovery period (P.Lokani, Department of Fisheries and
Marine Resources, PNG, pers. comm.).
The estimates of fisheries production in Tables 26 and 27 are the most accurate that can be obtained at present and are a “snapshot” of fisheries production in the
region towards the end of the 20th century, following 50 years of assisted development. Some countries, such as Fiji and the American territories, have very well
developed monitoring and survey programmes for estimating commercial fish production. However, in almost all countries, the estimates of subsistence fisheries
production must be computed empirically. This has been accomplished from dietary data where information has been recorded directly on per capita consumption of
fish or, more commonly, on the frequency of fish and shellfish consumption. Such information is collected during national nutritional surveys and epidemiological surveys
periodically carried out by government health departments. Such approaches are likely to be the only practical and costeffective method of determining subsistence
catches.
Advances have been made in the culture of commercial sessile invertebrates but other than the potential indicated by the successful deployment of spat collectors for
pearl oysters in the Cook Islands and French Polynesia, there are no demonstrable examples of successful replenishment of depleted populations by maricultured
stocks. This does not necessarily mean that all experiments have failed, but that the stock assessments used to date have not been sensitive enough to detect differences
between natural and “enhanced” recruitment, and this was the main reason why Palau has discontinued trochus reseeding experiments (N.Idechong, Marine Resources
Division, Palau, pers. comm.). Further research and development is required to determine if reseeding is feasible and costeffective, especially for those species that
do not have particularly low or erratic recruitment. It is generally accepted that recruitment is a major bottleneck in the rehabilitation of endangered giant clam
populations, but resources like trochus are much less fragile, and replenishment is probably best accomplished through fishery management. Such pure research may be
outside the financial and manpower resource priorities of most Pacific island fisheries departments and be more appropriately carried out by universities.
Page 508
Hatchery reef reseeding experiments may even contribute to unsustainable harvesting by suggesting that any current damage to stocks can be quickly repaired in the
future. It should be recognized that, at present, there is no universal panacea for overfishing (except the difficult decision to reduce either effort or catch). However, it
has been noted (M.Amos, Fisheries Department, Vanuatu, pers. comm.) that an active hatcherybased reseeding programme can have positive sociological benefits
when persuading coastal communities to place controls on fishing, particularly if that control is in the form of a moratorium and the “reseeded” organisms provide a
reason for voluntarily avoiding fishing in a particular area.
Management and stock conservation research in the Pacific islands should continue to be focused on mechanisms of limiting total harvest volume in socially and
economically feasible ways. The “traditional” approach to fisheries management that has evolved in the developed countries of the northern hemisphere involves a
relatively large fisheries research and management administration containing scientists, technicians, mathematicians, fisheries managers plus all the administrative support
staff that this entails. Sampling of landings at most ports is routinely conducted by the scientific staff, samples are taken from specimens for various studies, thousands of
otoliths are collected for ageing and research cruises are conducted to collect unbiased estimates of CPUE and to collect further catch samples. Standing stock
estimates are routinely computed from the biological data and the size of the catch for the following year or season is then estimated. The catch may then be divided up
between different countries based on whether the stock straddles several international boundaries or on political agreements such as the European Union Fisheries
Policy.
By contrast, most Pacific islands have small fisheries administrations, with little or no research capacity with which to gather information required for management.
Every village in a Pacific island is a potential landing site although increasing urbanization does lead to greater centralization of commercial landings. The archipelagic
nature of most countries, some stretching over hundreds of kilometres of ocean, and the relatively high costs of travel mean that those fisheries scientists and technical
officers that are available are restricted in their ability to collect information, most often to the capital city and to other major urban centres. An increasing number of
Pacific islanders are obtaining degrees in fisheries and marine sciences but there are still very few fisheries scientists in the region, particularly in the independent states,
where graduates of all disciplines are sought for higher government administration posts, or where salary levels are far more attractive in Pacific rim than Pacific island
countries. External funding is sometimes available to employ expatriate scientists and technical officers, but when they return to their home countries their experience,
and in some cases the information, leaves with them. The budgets available to South Pacific islands fisheries administrations are often marginal and usually insufficient for
necessary equipment. Even simple observations such as counting annuli in fish otoliths require some dissecting equipment and a low magnification compound
microscope costing several hundred dollars. These shortages of qualified and experienced fisheries staff coupled with limited financial resources mean that management
of coastal fisheries in much of the South Pacific is based on intuition rather than on collected observations and experience. Individuals can thus have enormous influence,
for good or for ill, on the management of resources over large groups of islands.
Johannes (1994) has argued that this very problem is an opportunity to seek a new paradigm for fisheries management in the South Pacific—one that is not based on
the conventional approach of intensive data gathering and analysis, but based on selfreinforcing feedback systems at the local level. Johannes argues that the very
limitations evident in most island fisheries departments are sufficient to make the conventional approach invalid as it is usually impossibly costinefficient to collect
sufficient data for the results to conform to the
Page 509
usual tests of significance. Instead, another approach is required where less emphasis is placed on “hard” data and more on gathering information from the fishermen
who prosecute the fishery. Johannes (1994) emphasizes that dataless management is not informationless management, but rather a mechanism that pays greater
attention to the information provided by fishermen on their assessment of a stock, and of the ways in which traditional and community measures were brought into play
to maintain stocks. Furthermore, such an approach would permit experimentation to assess the usefulness of given measures. For example, fishing on particular reefs is
routinely proscribed for a year in a certain Pacific island when a high chief dies, both as a mark of respect and to let marine resources build up for the memorial feast.
Such occasions permit observations on the effectiveness of reef closures. Johannes also notes that conventional fisheries research requires data collection over long time
periods before it can generate significant answers, whereas management decisions are usually required immediately, and that a trial and error approach, where errors
are part of the learning process, would be more suitable in the Pacific islands context.
It is likely that management of coastal fisheries in the South Pacific will develop into an amalgam of conventional approaches, including those where data are gathered
according to sampling criteria to test hypotheses, and the less data intensive approach advocated by Johannes (1994). There also needs to be greater attention given to
simply describing coastal fisheries to make better comparisons within and among countries that may, in turn, give greater insights into which management approaches
may work with a given fishery. The task of assembling the information for this paper revealed that there is a great deal of documented information on Pacific island
fisheries but much of this is in the informal or grey literature category and not readily available to fisheries scientists in the region. There are also a large number of
complete unknowns, as might be expected from a group consisting of thousands of islands, using hundreds of languages, scattered over a significant proportion of the
Earth’s surface.
For the large archipelagos of Melanesia, management of coastal fisheries, other than those for commercial export invertebrates, is not likely to become a pressing
issue in the near future. Management intervention on some of the smaller islands and atolls of Micronesia and Polynesia, however, appears to be immediately necessary
or will be required in the near future as populations continue to increase. Outright collapses of coastal fish stocks may be unlikely but, as stated earlier, some of the
larger slower growing species can become very vulnerable to modern gears and can be fished to extinction. The challenge for coastal fisheries management into the next
century will be to maintain production from the coastal zone as both human populations and the number of people turning to fishing as a livelihood increase.
Acknowledgements
This work was made possible through the generous support of the Overseas Development Administration of the United Kingdom to the South Pacific Commission’s
Integrated Coastal Fisheries Management Project and to N.V.C.Polunin. We thank all our colleagues who commented on the manuscript and who provided
information on various aspects of coastal fisheries in the South Pacific.
Page 510
Appendix 1. Summary of annual nominal coastal fisheries production by volume and value.
Fisheries statistics American Cook Federated States of Fiji French Guam Kiribati Marshall Nauru New Niue
Samoa Islands Micronesia Polynesia Islands Caledonia
Nominal coastal fisheries 267 982 6880 23253.04 6042.6 590.58 12314 2369 376.46 3480.5 115.4
production (t)
Nominal value of coastal 993000 3362444 12720944 64107438 28840189 2392045 18143667 3817717 848205 12968650 526224
fisheries production (US$)
Commercial fisheries production 51.6 124 637 6653.04 2351.6 118.18 3240 369 278.6 980.5 12
(t)
Nominal value of commercial 178762 314761 1483544 18340043 14371469 456413 4770000 714504 628605 3968650 54720
production (US$)
Subsistence fisheries production 215.4 858 6243 16600 3691 472.4 9084 2000 97.86 2500 103.4
(t)
Nominal value of subsistence 814238 3047638 11237400 45767395 14468720 1935632 13373667 3103213 219600 9000000 471504
fisheries production (US$)
Details of commercial fisheries production
Finfish
Reef & deepslope fish (t) 8.2 1 321 2917 909 9.5 1746 173 70.8 567.8 3.8
Reef & deepslope fish (US$) 32800 3552 577800 8527578 5454000 58378 2570000 275784 129885 1,769000 17182
Pelagic species (t) 43.1 24 198.5 955 916.5 109 690 13 207.8 3030.7 8.2
Pelagic species (US$) 144200 85526 357300 2654335 5787718 397515 1015833 34836 498720 255800 37538
Estuarine species (t) 708
Estuarine species (US$) 1457745
Invertebrates
Crustaceans (t) 0.3 20.5 436 3.1 0.08 13 34.1
Crustaceans (US$) 1762 196444 2693971 77500 520 19139 328700
Bêchesdemer (t) 334 77.7
Bêchesdemer (US$) 1998476 614650
Other echinoderms (t) 30.3
Other echinoderms (US$) 31087
Motherofpearl molluscs (t) 99 97 73 523 183 222.5
Motherofpearl molluscs (US$) 225683 352000 459638 3052251 403884 935000
Other molluscs (t) 1199 791 13.7
Other molluscs (US$) 517213 1165028 65500
Page 511
Fisheries statistics Northern Palau Papua New Pitcairn Solomon Tokelau Tonga Tuvalu Vanuatu Wallis & Western
Marianas Guinea Islands Futuna Samoa
Nominal coastal fisheries production 2966.2 1485.5 25554 8 11150 191 2362 927 2512 916.5 3488.5
(t)
Nominal value of coastal fisheries 12894231 4215251 63272908 16000 12749471 104509 4707849 755592 3467724 5422089 5389140
production (US$)
Commercial fisheries production (t) 141.2 735.5 4966 0 1150 0 1429 120 467 295.5 207.5
Nominal value of commercial 613804 2410059 22096908 0 4343811 0 2806641 97811 1514364 2316729 319066
production (US$)
Subsistence fisheries production (t) 2825.0 750 20,588 8 10,000 191 933 807 2,045 621 3,281
Nominal value of subsistence fisheries 12280427 1805192 41176000 16000 8405660 104509 1901208 657781 1953360 3105360 5070074
production (US$)
Details of commercial fisheries production
Finfish
Reef & deepslope fish (t) 65.6 492.1 1,100 87 1,254 77 274 165.6 171.7
Reef & deepslope fish (US$) 295353 1184526 2200200 73129 2399094 62762 453396 910800 263745
Pelagic species (t) 74.6 630 150 43 110.00 15.3
Pelagic species (US$) 309750 1260000 305660 35049 825000 23643
Estuarine species (t) 908
Estuarine species (US$) 1960000
Invertebrates
Crustaceans (t) 1 14.4 1,240 2 25 26 3.4 20.5
Crustaceans (US$) 8701 125533 10500000 101887 127358 37400 31678
Bêchesdemer (t) 546 622 24
Bêchesdemer (US$)
Other echinoderms (t)
Other echinoderms (US$)
Motherofpearl molluscs (t) 229.0 470 439 143 16.5
Motherofpearl molluscs (US$) 1100000 1400000 1675818 847342 543529
Other molluscs (t)
Other molluscs (US$)
Page 512
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AUTHOR INDEX
© A.D.Ansell, R.N.Gibson and Margaret Barnes, Editors
UCL Press
References to complete articles are given in bold type; references to page numbers are given in normal type; references to bibliographical lists are given in italics.
Abbott, D.P. See Haderlie, E.C., 167; 226

See Newman, W.A., 306, 359, 361, 381, 382; 391
Abbott, D.T., 381; 384
See Mix, M.C., 391
Abbott, R.T., 182; 222
Abele, L.G. See Coen, L.D., 154
AbeleOeschger, D., 288; 293
Abeyasekera, S., 401; 512
Achituv, T. See NeubergerCywiak, L., 228
Achituv, Y., 351, 352, 353; 384
Acrivos, A., 78; 101
Acrivos, A. See Frankel, A.N., 88; 101
Adams, T. See Nash, W., 524
Adams, T.J. H., 404, 413, 431, 444, 472, 473, 474, 475, 478, 480, 482, 483, 484, 485, 494, 496, 497, 501, 503; 512
See Dalzell, P., 395–531
See Lewis, A.D., 522
See Sharma, S.P., 459, 460; 528
Adey, W.H. See Brawley, S.H., 123; 153
Aggarwal, B.M. See Anand, S.P., 67
Ahsanullah, M. See Newell, R.C., 298
Aini, J. See Hair, C., 490, 496; 520
Aitaoto, F. See Craig, P., 516
Alagarswami, K., 205, 206; 222
Alarcon, E., 192; 222
Alberte, R.S. See Kirchman, D.L., 157
See Mazzella, L., 117, 118, 119, 133, 149; 158
Aldan, C. See Adams, T.J.H., 572
Aleem, A.A., 198; 222
Alerstam, T., 253, 291; 254, 293
Alexander, S.K., 282, 283; 293
Alfred, A. See Sims, N.H., 528
Alfred, V. See Adams, T.J. H., 512
AlHamandani, E. See Moyse, J., 259
Alkire, W.H., 428; 512
Alldredge, A.L., 139; 153
Alldridge, A.E. See Richardson, J.R., 230
Allen, K.R., 441; 512
Almeida, P., 180; 222
Ames, J. See Wendell, F., 232
Amesbury, S.S., 407, 410, 413, 415, 421, 446, 449, 450, 453, 459, 479, 488; 512
Amon, M.W., 81; 101
Amos, M. See Adams, T.J. H., 512
See Nash, W., 524
Amouroux, J.M., 195; 222
Anand, S.P., 11,23,24,27; 66
Anderson, D.T., 250, 304, 320, 321, 322, 323, 324, 326, 327, 331, 334, 339, 342, 343, 348, 353, 354, 366, 368, 369, 373; 254, 384
See Walker, G., 332; 393
Anderson, T.A. See Braley, H., 293
Ando, T., 195; 222
Anell, B., 405; 512
Anonymous, 2, 7, 8, 10, 56, 397, 399, 401, 403, 408, 410, 411, 415, 418, 419, 420, 421, 431, 434, 438, 446, 449, 450, 452, 459, 467, 470, 472, 473, 474, 479, 484, 486, 488, 490, 491, 492, 497, 498, 504,
505; 66, 512, 513
Ansari, Z.A. See Parulekar, A, H., 229
Ansell, A.D., 183, 195, 196, 197, 201, 204, 205; 222
See McLachlan, A., 163–232
Antonio, R.S. See Hamm, D. C, 520
Antony, M.K., 11; 66
See Rao, K.H., 69
Appleton, R.D., 241, 242, 253; 254
Ardizzoni, S.C. See Hamilton, P.V., 295
Ardré, F., 308, 310; 384
AriasGonzalez, J.E., 423, 425; 513
Arntz, W.E., 192, 193; 222
ArreguínSánchez, F. See Defeo, O., 224
Arrontes, J. See Viejo, R.M., 287; 301
Arsuffi, T.L. See Barlocher, F., 293
Arthur, W. See Clarke, B., 255
Ash, K. See MatengaSmith, T., 523
Ash, V.B., 242; 254
Atkinson, M.J., 430; 514
See Grigg, R.W., 519
Atkinson, W.D., 237, 238, 250; 254
Austin, A.P., 369; 385
Avila, M. See Santelices, B., 299
Ayling, A.M. See Zann, L.P., 484; 531
Ayres, D.L., 173, 174, 175, 176, 177, 178, 220; 222
Azevedo, C., 367; 385
Babcock. J. See MatengaSmith, T., 523

Babin, M. See Amesbury, S.S., 449, 450; 512
Babu, M.T., 13, 33, 55, 56, 59, 60; 66
See Murty, V.S. N., 68
See Sastry, J.S., 69
Page 534
See Suryanarayana, A., 70

Babu, S. V. See Rao, R.R., 69
Bach, C., 282; 293
Bach, F. See MatengaSmith, T., 523
Bach, J. See Elymore, J., 518
Backeljau, T., 236; 254
Badcock, J. See Coyne, T., 516
See Elymore, J., 518
Badino, G., 195; 222
Bagnis, R., 423, 453; 514
Bahulayan, N., 56, 61; 66, 70
See Unnikrishnan, A.S., 61; 70
Baillon, N., 424; 514
Baker, M.A., 95; 101
Bakun, A., 21; 66
Balaramamurty C. See Ramasastry, A.A., 10; 68
Balaramamurty, C.B., 10, 11, 25, 56; 66
Ballantine, D.L., 115, 117; 153
Ballantine, W.J., 271, 272, 273; 293
Ballment, E. See Benzie, J.A.H., 486; 514
Bally, R., 199; 222
Bandel, K., 237; 254
Banta, W.C. See Stanhope, H.S., 299
Baquie, B., 407, 410; 514
Bargibant, G. See Kulbicki, M., 521
Barker, K.M., 272, 273, 274, 275, 276, 279, 286, 287, 289; 293
Barlocher, F., 280, 283; 293
Barnes, H., 304, 306, 308, 309, 310, 321, 324, 326, 328, 329, 330, 335, 337, 338, 339, 340, 341, 342, 344, 345, 348, 349, 353, 355, 359, 361, 363, 364, 366, 367, 368, 370, 375, 376, 378, 380; 385
See Achituv, Y., 351, 352, 353; 384
See Klepal, W., 373, 374, 375; 390
See Morris, R.J., 350, 351, 352; 391
See Munn, E.A., 367; 391
Barnes, M., 303–94 362, 366, 368, 374; 385
See Barnes, H., 304, 308, 310, 330, 344, 348, 349, 364, 365, 366, 370, 374, 375, 376, 378; 385
See Klepal, W., 390
Barnett, P.R.O. See Ansell, A.D., 222
Barsdate, R.J. See McRoy, C.P., 158
Bastida, R., 190; 223
Batchelor, G.K., 86, 88, 89, 90, 92, 93, 94, 96, 97, 100; 101
Bate, G.C. See McLachlan, A., 201; 227
See Talbott, M.M.B., 261
Batham, E.J., 323, 324, 334, 361, 372; 385, 386
Batty, M., 465; 514
Bauer, T.R., 139; 153
Baumgartner, A., 6, 7, 8; 66
Bayed, A., 195, 197; 223
See Guillou, J., 194, 195; 226
BaylissSmith, T., 410, 411; 514
Bayne, B. L., 121, 146, 276; 153, 293
Beardmore, J.A., 235, 244; 254
Beauchamp, P.de, 305, 307, 308, 310; 386
Beaumont, A.R., 236; 254
Bebout, B. See Kohlmeyer, J., 280, 283; 296
Beckey, B.B. See Hasse, J.J., 520
Beddington, J.R. See Rosenberg, A.A., 501; 527
Begon, M. See Hart, A., 251; 256
See Naylor, R., 238, 240, 245; 259
Behrens Yamada, S., 246, 277, 279, 286; 254, 255, 293
Beklemishev, C.W., 85; 101
Bell, J.D., 120, 122, 135, 147, 149; 153
See Richards, A.H., 527
Bell, J.S., 399, 431; 514
Bell, L. See Adams, T.J.H., 512
See King, M., 490, 493; 521
See Zann, L.P., 531
Bell, L.J. See Richards, A.H., 527
Bell, R.S., 494; 514
Bell, S.S. See Hall, M.O., 117, 148; 155
Benet, J. See Bagnis, R., 514
Benjamin, K.J. See Lethbridge, R.C., 157
Benner, R. See Amon, M.W., 81; 101
Bennett, I. See Dakin, S.M., 206, 207; 224
Bennette, E.B., 53, 54, 55; 66
Benson, M.R. See Norton, T.A., 143, 290; 159, 298
Benzie, J.A.H., 486; 514
Berdálet, E., 97; 101
Berg, H.C., 71, 72, 78, 79, 82, 83, 107; 101
Berger, E., 236, 248; 255
Berger, E.M., 246; 255
Bergerard, J., 252; 255
See Caugant, D., 252; 255
Berman, J., 279; 293
Bernard, F.R., 305, 307, 321, 358, 359, 361, 379, 381, 382, 383; 386
Berry, A.J., 247, 251; 255
Bertness, M.D., 272, 273, 274, 276, 278, 279, 285; 293
See Kemp, P., 242; 258
Bertram, I. See Adams, T.J.H., 512
Beu, A.G., 207; 223
Beverly, S., 446, 452; 514
Beverton, R.J.H., 496, 503; 514
Bhattathiri, P.M.A. See Rhadakrishna, K., 68
Bhavanarayana, P.V. See Ganapathi, P.N., 67
Bianchi, G. See Fischer, W., 453; 225, 518
Bird, J.B., 271; 293
Bishop, M.W.H., 307, 310; 386
BishopLowen, B. See Bourne, N., 223
Blaber, S.J.M., 404, 410, 420, 460, 466; 514
See Milton, D.A., 523
See Rawlinson, N.J.F., 527
Black, R., 267, 268; 293
See Johnson, M.S., 248; 258
Blackstock, J. See Barnes, H., 321, 375, 376, 378; 385
Blackstone, N.W., 278; 293
Blair, D.F. See Parkinson, J.S., 79; 103
Blanc, M., 481, 482; 514
Bleil, D.F., 281; 293
Page 535
Blum, H.F. See Florkin, M., 329; 388

Blundon, J.A., 282; 293
Bodoy, A., 195, 197; 223
See Ansell, A.D., 195, 196; 222
See Plante Cuny, M.R., 197; 229
Boekschoten, G.J. See Juch, P.J.W., 272; 296
Bolivar, I., 307, 310; 386
Bolton, L.A., 479, 481; 514
Bone, D. See Pauls, S.M., 229
Boolootian, R.A. See Leighton, D., 237; 255
Boonruang, P. See Sundstrom, B., 70
Borjesson, D.L., 282; 293
Borowitzka, M.A., 111, 114, 115, 116, 117, 118, 119, 120, 132, 133, 134, 135, 141; 153
See Lethbridge, R.C., 157
See Reichelt, J.L., 118; 159
Borum, J., 117, 118, 122, 127, 132, 133, 141, 142, 143, 150; 153
See SandJensen, K., 118, 135; 160
Boss, E. See KarpBoss, L., 71107
Boulding, E.G., 236, 240, 241, 242, 252; 255
Bour, W., 483; 514
Bourget, E., 353; 386
Bourne, N., 172, 174, 177; 223
See Quayle, D.B., 174; 230
Bouvier, E.L., 308; 386
Bowen, J.D., 95; 101
Boy, R.L., 445; 514
Brainerd, K.E., 92, 95; 101
Braley, H., 270; 293
Braley, R.D., 405, 467, 484, 488, 491, 493, 496; 514
Bramwell, M.D., 116, 117, 133, 134; 153
Branch, G.M., 267, 268; 293
See McQuaid, C.D., 292; 297
Branch, M.L. See Branch, G.M., 267, 268; 293
Brandt, E., 323; 386
Brandwood, A., 240; 255
Branford, J.R., 495; 514
Brauner, J.F., 135; 153
Brawley, S.H., 109, 119, 123; 153
Brearley, A., 122; 153
See Jernakoff, P., 109–62
Breeman, A.M., 288; 293
Breese, W.P., 172; 223
Bremec, C. See Bastida, R., 223
Brenchley, G.A., 272, 273, 277, 278; 294
Brennen, C., 83; 101
Brenner, H., 84; 101
See Happel, J., 85; 102
Brey, T. See Arntz, W.E., 222
Brito, P.J., 180, 181, 182; 223
Brittan, C. See Cambridge, M.L., 154
Brix, H., 117; 153
Broch, H., 304, 306, 308, 312, 313, 317, 318, 319, 320, 343, 354, 357, 378; 386
Brock, V.E., 426; 514
Brostoff, W.N., 119; 153
Brouard, F., 437, 439, 440; 514
Brouns, J.J.W.M., 114, 133, 135; 153
Brown, A.C., 200, 201; 223
See Matthews, S., 227
See StentonDozey, J.M.E., 201; 231
See Trueman, E.R., 201; 231
Brown, A.F. See Bertness, M.D., 293
Brown, I.W., 414, 423, 425, 487, 489, 494, 496, 505; 515
Bruijsten, J. See Kronig, R., 82; 102
Brumley, B.H., 96; 101
Bruntz, L., 334; 386
Bryan, G.W., 264; 294
Buat, P. See Poupin, J., 525
Buckeridge, J.S. , 304; 386
See Foster, B.A., 304, 305; 388
BucklandNicks, J. See Boulding, E.G., 255
Buckley, R., 449; 515
Buckley, T. See Buckley, R., 515
Buia, M.C. See Mazzella, L., 158
Bukurrou. A. See Adams, T.J.H., 512
Bull, C.M. See Chilton, N.B., 269; 294
Bulleid, N.C. See Tranter, D.J., 161
Bulthuis, D.A., 111, 114, 117, 118, 133, 134, 135; 153, 154
See Shepherd, S.A. 160
Bunting D.L. See Mix, M.C., 391
Burchmore, J.J. See Bell, J.D., 153
Burge, R. See Wendell, F., 232
Burnett, B.R., 327, 328, 331, 333, 334, 340, 349; 386
Burollet, P.A., 308; 386
Burt, G.R. See Bryan, G.W., 294
Burton, R., 245; 255
Buschmann, A.H., 123, 146; 154
Butler, A.J., 482; 515
Byers, B.A., 238; 255
Bynum, K.H. See Williams, A.B., 138; 161
Cabañas Ruesgas, F. See Ardré, F., 384

Cabot, E.L. See Smith, B.D., 299
Caddy, J.F., 220; 223
Caillart, B., 410, 412, 415, 419, 423, 424; 515
See Morize, E., 425; 524
Caine, E.A., 123, 124, 150; 154
Cairns, K.D. See Nelson, W.G., 158
Caldwell, J. See Gaines, M.S., 256
Callahan, R.A. See Shokes, R., 334; 393
Calow, P., 250; 255
Calvin, J. See Ricketts, E.F., 392
Cambridge, M.L., 111, 114, 118, 120, 121, 132, 150; 154
See McComb, A.J., 158
Campana, S.E., 502; 515
Campbell, E.E. See Talbott, M.M.B., 261
Campbell, R. See Tranter, D.J., 161
Camus, P.A. See Castilla, J.C., 193; 223
Canelli, E., 85; 101
Page 536
Cannon, H.G., 322, 327, 328, 339, 368; 386

Capezzani, D. See Olivier, S., 229
Carbonara, P.A. See Virnstein, R.W., 116; 161
Carcelles, A., 189; 223
Cardosa, A.C., 361, 362, 366; 386
Carey Jr, A.G. See Hogue, E., 173; 226
Carlos, G. See Samoilys, M., 426; 527
Carlot, A. See Schaan, O., 528
Carlot, A.H., 434, 440; 515
Carlson, S.S. See Wilson, A.C., 262
Carlton, J.T., 271; 294
See Berman, J., 279; 293
See Brenchley, G.A., 272, 273, 277, 278; 294
Carmichael, A. See Bourne, N., 177
Carreto, J. See Olivier, S., 229
Carriquiriborde, L., 168; 223
Cary, J.L. See Masini, R.J., 158
Cascudo, J. See Defeo, O., 224
Casola, E., 133; 154
Cassie, R.M., 208, 209, 210, 211; 223
Castaño, J.T., 322; 386
Castenholz, R.W., 265; 294
Castilla, J.C., 193, 220 ; 223
See Defeo, O., 224
Cattaneo, A., 145; 154
Caugant, D., 252; 255
See Bergerard, J., 252; 255
Caziot, Le Commandant, 308; 386
Ceccaldi, H.J. See Ando, T., 195
Cernhorsky, W.O., 404; 515
Chaffee, C., 317; 386
Chaffee, J., 353, 355; 386
Chakraborty, R. See Nei, M., 259
Chambers, A., 410, 463; 515
Chan, A.T. See Harrison, P.G., 118; 155
Channells, P.W. See Bell, R.S., 514
Chapau, M. See Haines, A., 445, 451; 519
See Lokani, P., 477, 482; 522
Chapau, M.R., 410, 418, 421, 423, 428, 434, 438, 464, 465, 467, 470, 471, 481, 488, 492, 493, 494, 502; 515
See Wright, A., 521
Chapman, A.R.O., 287, 288, 291; 294
See Barker, K.M., 272, 273, 274, 275, 276, 279, 286, 287, 289; 293
See Denton, A.B., 294
See McCook, L.J., 275, 288, 289, 292; 297
See Parker, T., 298
Chapman, L.B., 405, 420, 445, 446, 449; 515
See Preston, G.L., 526
Chapman, M.G., 268, 292; 294
See Underwood, A.J., 268; 300
Chari, S.T., 206; 223
Charlton, L. See Borowitzka, M.A., 153
Chassé, C., 195; 223
Chatterji, A. See Parulekar, A.H., 229
Chauvet, C., 487, 490, 491, 493, 494; 515
See Grandin, P., 487, 490, 491, 493, 494; 519
Chen, C. See Lewin, J., 227
Chesher, R.H., 483; 516
Chessa, L.A. See Russo, G.F., 159
Chessel, D. See Mouëza, M., 194; 228
Chew, K.K. See Schink, T.D., 230
Chia, F.S., 249; 255
See Lewis, C.A., 359, 360, 361, 362, 363, 364, 365, 366, 368; 390
See Martel, A., 246, 249, 290; 259, 297
Chiffings, A.W. See Cambridge, M.L., 154
See Silberstein, A.K., 160
Chilton, N.B., 269; 294
Chisholm, S.W. See Bowen, J.D., 101
Choi, H.Y. See Rohsenow, W.M., 77; 103
Chow, V., See Mastro, E., 259
Choy, S., 405, 488, 491, 493, 495; 576
Christensen, J.T. See Hylleberg, J., 240, 265, 272; 257, 296
Christian, E. See Bagnis, R., 514
Christiansen, A. See GeertzHansen, O., 155
Christiansen, H. See Olivier, S., 229
Church, M. See South, G.R., 529
Chwang, A.T., 83; 101
Cillauren, E., 449; 516
See David, G., 410, 411, 428, 466, 498; 517
Cimberg, R.L., 361, 362, 363, 364; 387
Cinelli, F., 132, 134, 141; 154
Clare, A.S., 326, 327; 387
Clarke, A.H., 271; 294
Clarke, B., 238, 243; 255
Clarke, B.C., 243; 255
Clarke, D.J. See Donn Jr, T.E., 225
Clarke, R., 491, 493; 516
Clarke, S.M. See Sergeev, V.N., 160
Clarke, T.A., 460; 516
Clarner, J.P. See Tenore, K.R., 161
Clift, R., 78, 79, 84, 85, 90; 101
Coates, D., 464, 466, 467, 471, 490, 492; 516
See Grassland, S., 256
Cockcroft, A.C., 201; 223
See du Preez, H.H., 207
Coe, W.R., 165, 166, 168, 169, 173; 223
Coen, L.D., 148; 154
Coetzee, P. See McLachlan, A., 227
Colborn, J.G., 23, 24, 27, 34; 67
Cole, T.J., 237; 255
Coles, R., 401; 516
Collett, L.C., 137; 154
See May, V., 158
Collette, B.B., 404, 466; 516
Collins, A.J. See Collett, L.C., 154
See May, V., 158
Colmenares, R. See Penchaszadeh, P.E., 229
Comfort, A., 237; 255
Comps, M., 196; 223
Conacher, M.J., 120; 154
Conand, C., 473, 475, 476; 516
Page 537
Conand, F., 459, 460; 576

Confer, D.R., 72; 101
Connor, M.S., 278; 294
Cook, I.H. See Kirkman, H., 114; 157
Cook. J., 451, 477; 516
Cook, L.M., 238, 239, 240; 255
Cooke, A., 432; 516
Copland, J.W., 483; 516
See Blaber, S.J.M., 460; 514
Cormaci, M. See Cinelli, F., 154
Cornwall, I.E., 306, 313, 317, 323, 324, 381; 387
Corral, L. See Azevedo, C., 367; 385
Correa, J. See Santelices, B., 288; 299
Coscarón, S., 187; 223
Coutures, E., 424, 502; 516
Coyne, T., 409; 516
Craig, M.P. See Polovina, J.J., 525
Craig, P., 410, 412, 449, 498; 516
Crane, P. See Coates, D., 516
Cranfield, H.J., 214, 215, 216; 223, 224
Crank, J., 73; 101
Crean, K., 473, 474, 475; 516
Crecelius, E.A. See Drum, A.S., 225
Creese, R.G., 237; 255
See Underwood, A.J., 237; 262
Crenshaw, D.G., 333; 387
Crisp, D. See Foltz, D.W., 256
Crisp, D.J., 244, 246, 248, 272, 307, 310; 256, 294, 387
See Bishop, M.W.H., 386
Crocombe, R., 505; 517
Crossland, S., 236; 256
Cruz, J. See Adams, T.J.H., 512
Cruz, T., 308, 359, 360, 383; 387
Csanady, G.T., 18; 67
Cubit, J. See Lubchenco, J., 286; 297
Cubit, J.D., 269; 294
Culver, M.E., 83; 101
Curran, M.C. See Virnstein, R.W., 117; 161
Curray, J.R. See Kolla, V., 67
Currey, J.D., 242; 256
See Cook, L.M., 255
Currie, R., 1; 67
Cusack, P., 407, 445; 517
See Beverly, S., 446, 452; 514
See Chapman, L.B., 405, 420, 445, 446, 449; 515
Cusack, P. See Taumaia, P., 410, 418, 449, 457; 529
Cusack, P. See Watt, P., 452; 530
Cushing, F.A. See Amesbury, S.S., 512
Cussler, E.L., 77, 82; 101
Cutler, A.N., 56; 67
Daborn, G.R. See Strong, K.W., 143; 160

Dabrowski, H. See Estep, M.F., 287; 294
Dade, W.B. See Jumars, P.A., 102
Dahlstrom, W.A. See Miller, D.J., 228
Dakin, S.M., 206, 207; 224
Dalzell, P., 395–531; 406, 410, 411, 414, 415, 417, 418, 419, 420, 421, 422, 423, 428, 432, 434, 435, 438, 439, 440, 443, 446, 447, 449, 452, 453, 454, 455, 456, 457, 458, 460, 461, 462, 474, 477, 481, 484,
488, 490, 492, 498, 502, 503, 504; 517
See Adams, T.J.H., 484, 494, 496, 497, 501; 512
See Chapau, M.R., 434; 515
See Diplock, J.H., 418, 439; 518
See Kitalong, A., 410, 415, 423, 424, 425, 428; 521
See Patiale, H., 410, 418, 498; 525
See Smith, A., 406, 407, 410, 412, 419, 421, 426, 427, 504; 528
See Wright, A., 531
Daniel, A., 307; 387
D’Antonio, C., 285; 294
Darwin, C., 303, 304, 305, 306, 307, 308, 311, 312, 313, 318, 321, 322, 326, 327, 334, 339, 354, 355, 366, 368, 373, 382; 387
Das, P.K. See Varadachari, V.V.R., 70
Dashwood. J., 486; 517
See Adams, T.J.H., 480; 512
David, G., 410, 411, 428, 466, 498; 517
Davies, M.S., 272; 294
Davies, P.A. See Sigurdsson, J.B., 231
Davis, G.E. See Dugan, J.E., 220; 225
Dawson, E.W., 215; 224
Dayton, P.K., 306, 308, 366; 387
See Tegner, M.J., 142; 160
de Alava, A., 189, 218; 224
See Defeo, O., 188, 218, 220; 224
Debao, A. See Dalzell, P., 410, 414, 418, 419, 421, 422, 428, 446, 447, 449, 481, 498, 504; 517
de Coo, A. See Fernandez, J., 225
Dee, A.J., 425; 517
Defant, A., 54; 67
Defeo, O., 169, 186, 187, 188, 189, 190, 191, 193, 194, 218, 220; 224
See de Alava, A., 189; 224
See Masello, A., 187; 227
See McLachlan, A., 163–232; 227
Degiovanni, C., 195, 197; 224
Delathiere, S., 448, 492, 494, 495; 517
Delaune, J.M., 488, 491; 517
De Mahieu, G., 180, 182, 184; 224
See Penchaszadeh, P. E., 229
DeMartini, E. See Polovina, J.J., 525
DeMartini, E.E. See Ellis, D.M., 441; 518
Deming, J.W. See Jumars, P.A., 102
de Moreno, J.A. See Olivier, S., 229
denHartog, C., 111; 154
See Larkum, A.W.D., 111; 157
Denley, E.J. See Underwood, A.J., 300
Dennison, W.C., 111; 154
Denoux, G.J. See Shirley, T.C., 299
Denton, A.B., 274; 294
De RooijSchuiling, L.A. See Beu, A.G., 207; 223
de Ruyck, A. See McLachlan, A., 227
Page 538
DeslousPaoli, J.M. See Sornin, J.M., 160

DeSouza, S.N., 11; 67
Dethier, M.N. 285, 286; 294
Dethier, M.W. See Steneck, R.S., 160
Dettmer, J.W. See Logan, E.L., 72, 91; 102
Devassy, V.P. See Rhadakrishna, K., 68
de Villiers, G., 199, 202, 203; 224
Diaz, M.C., 184; 225
Dickie, L.M. See Mallet, A.L., 258
Dickinson, R.E., 494; 518
Diefenbach, T. See Martel, A., 249; 259
Dignan, C. See MatengaSmith, T., 523
Diplock, J.H., 418, 439; 518
Dippolito, A., 277; 294
Dix, T.L., 282; 294
Doering, P.H., 146; 154
Doering, V. See Diaz, M.C., 154, 184
Doherty, P. See Brown, I.W., 515
See Kulbicki, M., 521
Doherty, P.J., 503; 518
See Meekan, M., 259
Donn, T.E., 200, 201, 202; 225
Donn Jr, T.E., 199, 200, 202, 217; 225
Doonan, I.J. See Cranfield, H.J., 224
Dorrepaal, J.M., 85; 101
DorstHansen, W.See Korson, L., 102
Dortch, Q., 85; 101
Double, M. See Grahame J., 256
Douglas, G., 399; 518
Doumergue, F., 382; 387
Dowdell, F. See Hamm, D. C., 520
Dowidar, N.M., 195, 197, 198; 225
Doyle, R.W. See Newkirk, G. F., 236, 240; 259
Doyle, R.W. See Shaddock, P.F., 144, 149; 160
Drew, A.W., 424; 518
Drew, K.M., 378; 387
Drum, A.S., 178; 225
Ducker, S.C., 116, 132, 135; 154
Dudley, D.L., 281; 294
Dudley, P.L., 366, 367, 369; 387
Dudley, R., 242; 256
Duffy, J.E., 142, 143; 154
See Hay, M.E., 155
Dugan, J. See McLachlan, A., 227
Dugan, J.E., 167, 200, 220; 225
Dugan, P.J., 138, 139, 140; 154
Düing, W., 57; 67
duPreez, H.H., 201; 225
Durance, C. See Harrison, P.G., 118; 155
du Toit, P. See Donn Jr, T.E., 225
Dye, A.H. See McLachlan, A., 227
Eastman, R.C., 372; 387

Ebert, T.A., 476, 491, 493, 494; 518
Eckman, J.E. See Lewin, J., 227
Edgar, G.J., 119, 122, 123, 135, 136, 137, 138, 139, 140, 141, 147, 148, 149; 154, 155
See Howard, R.K., 156
Efford, I.E., 219; 225
Eldredge, L., 497; 518
Elliott, J.A. See Oakey, N.S., 92; 103
Ellis, D.M., 441; 518
Ellis, T. See Poupin, J., 525
Ellway, C.P., 462; 518
ElNady, F.E. See Dowidar, N.M., 195, 197, 198; 225
Elner, R.W. See Vadas, R.L., 271, 272, 274, 279, 291; 301
Els, S.F. See Donn Jr, T.E., 200; 225
Elston, R.A., 173, 174; 225
See Drum, A.S., 225
Ely more, A., See Ely more, J., 518
Elymore, J., 449; 518
Emery, A.R., 410, 411, 419; 518
Emlen, J. See Spight, T.M., 250; 261
Endler, J.A., 238; 256
Ephremov, V.V. See Kartavtsev, Y.P., 247; 258
Eppley, R.W., 83; 101
Epstein, N. See Masliyah, J.H., 79, 85; 103
Erskine, J.S. See Clarke, A.H., 271; 294
Estep, M.F., 287; 294
Estes, J.A., 168, 173, 175, 176, 177; 225
Etchevers, S.L., 180, 181, 182, 184; 225
Etter, R.J., 239, 256
Euan, J. See Defeo, O., 224
Evans, C., 492, 495; 518
Evans, D., 458, 460; 518
Fairweather, P.G., 267, 268, 269; 295

See Jernakoff, P., 268; 296
See Moran, M.J., 297
Fairweather, P.J. See Underwood, A.J., 292; 300
FallerFritsch, R.J., 250; 256
Fancett, M.S., 138; 155
FAO, 402, 463, 498; 518
Farache, V., 182, 184, 185; 225
See Penchaszadeh, P.E., 229
Farias, A., See Carriquiriborde, L., 223
Farman, R. See Chauvet, C., 487, 490, 491, 493, 494; 515
Farrell, T.M. See Satchell, E.R., 358; 392
Faustino L.A. See Talavero, F., 206; 231
Feder, H.M. See Orth, F.L., 229
Fenchel, T., 84; 101
See Vadas, R.L., 161
Fenical, W. See Hay, M.E., 155
Fenwick, G.D., 148; 155
Fernandez, J., 197; 225
Ferreira, B. See Brown, I.W., 515
Ferreira, B.P., 425; 518
Ferrer, L., See Carriquiriborde, L., 225
Fevolden, S.E., 244; 256
Fielden, L.J. See Hodgson, A.N., 201; 226
Fielder, D.R., See Brown, I.W., 489, 496, 505; 515
Filún, L. See Jaramillo, E., 226
Page 539
Fischer, P., 308; 387

Fischer, W., 194, 453; 225, 518
FischerPiette, E., 307, 308, 310, 382; 387
See Ardré, F., 384
See Crisp, D.J., 307, 310; 387
Fish, J.D., 251; 256
Fisher, F.M. See Vaughn, C.C., 280, 281; 301
Fisher, K. See Black, R., 293
Fitch, J.E., 165, 166, 167, 168, 171; 225
See Coe, W.R., 165, 166, 167, 168, 169; 223
Fitt, W.K., 483; 518
Fitzgerald, W.J. See Tsuda, R.T., 530
Fitzpatrick, L.C. See Velez, A., 232
Flannery, B.P. See Press, W.H., 103
Fleming, R.H. See Sverdrup, H.U., 70
Fletcher, C.R. See Lowell, R.B., 258
Fletcher, W.J., 496; 518
Flint, E.N. See Polovina, J.J., 525
Flores, T. See Adams, T.J.H., 512
Florkin, M., 329; 388
Fluck, W.H., 182; 225
Foltz, D.W., 244; 256
See Liu, L.L., 258
See Zouros, E., 244; 262
Fomicev, A.V., 57; 67
Fonseca, M.S., 111; 155
Forcucci, D. See Dugan, J.E., 225
Ford, N.J. See Ducker, S.C., 154
Ford, R.F. See Ebert, T.A., 491, 493, 494; 518
Foreman, R.E. See Smith, B.D., 299
Forgacs, O.L., 95; 101
Forman, S.L. See Salvigsen, O., 299
Fortes, M.D. See Klumpp, D.W., 157
Foster, B.A. 304, 305, 340, 343, 354; 388
See Newman, W.A., 305; 391
Foster, M., 380; 388
Fotheringham, N., 282; 295
Fox, W.W., 429, 495; 518
Fralick, R.A.F., 289, 290; 295
Franc de Ferriere, M. See Caillart, B., 515
Frankel, A.N., 88; 101
Freeman, K.R. See Mallet, A.L., 258
Freeman, P.M. See Cook, L.M., 238; 255
Frenkiel, L., 196; 226
See Ansell, A.D., 222
See Mouëza, M., 197; 228
See Pichon, Y., 229
FrenkielRenault, L. See Mouëza, M., 195; 228
Fresi, E. See Casola, E., 154
See Russo, G.F., 159
Fretter, V., 236, 247, 289; 256, 295
Frey, H.W., 167, 169, 171; 226
Frid, C.L.J., 276, 277, 278; 295
Friedlander, S.K., 85; 101
Frost, P.G.H. See McQuaid, C.D., 297
Frusher, S.D., 488, 489, 490, 494, 495; 518, 519
Fry, B., 123; 155
See Kitting, C.L., 157
Fuerst, P.A. See Nei, M., 259
Fuhs, G. W. See Canelli, E., 85; 101
Furnari, G. See Cinelli, F., 154
Fusimalohi, T., 439; 519
Fyhn, H.J.., 328, 330, 331, 332, 347, 348, 349; 388
See Petersen, J.A., 392
Gaines, M.S., 247; 256

Gaines, S.D., 290; 295
See Lubchenko, J., 121, 266; 158, 297
Gallagher, J.F., 23, 41, 50, 52; 67
Gallardo, C.S., 250; 256
Galzin, R., 410, 414, 426, 428; 519
See AriasGonzalez, J.E., 513
Gamba, A.L. See De Mahieu, G., 180; 224
Gambi, M.C., 122, 123, 137; 155
See Mazzella, L., 158
Ganapathi, P.N., 10; 67
Garbett, S.D. See Cook, L.M., 239; 255
Gargett, A.E., 92; 102
Garner, S.P. See Fevolden, S.E., 244; 256
Garrault, H., 322, 333; 388
Garrett, C., 54; 67
GartnerKepkay, K.E. See Mallet, A.L., 258
Gastelu, A. See Carriquiriborde, L., 223
Gavis, J., 71, 83; 102
See Pasciak, W.J., 75, 91, 105; 103
Gayanilo, F.C., 192; 226
GeertzHansen, O., 150; 155
Geiselman, J.A., 279, 286; 295
Geldiay, R., 195; 226
Geller, J.B., 274, 276; 295
Gentle, M.T., 475; 519
George, C.J., 197; 226
George, R.W., 490; 519
Gerpe, M. See Bastida, R., 223
Gianuca, N.M., 187, 188, 190; 225
Gibbs, P.J. See Collett, L.C., 154
Gibson, C.H. See Baker, M.A., 95; 101
See Thomas, W.H., 97; 104
Gibson, R. See Zimmerman, R.J., 162
Gibson, R.N. See Ansell, A.D., 197; 222
Giesel, J.T. 309; 388
Gilfillan, E.S. See Trussell, G.C., 262
Gillet, B. 479; 519
Gillett, R., 406, 453, 455, 457; 519
Gillett, R.D., 421; 519
Gillilan, W., 168; 226
Gilly, W.F., 280; 295
GinaWhewell, L., 410, 481; 519
Goddard, J.D. See Acrivos, A., 78; 101
Godfrey, M.C.S., 207; 225
Goeden, G.B., 425; 519
See Mcpherson, G.R., 523
Goering, J.J., 118; 155
See McRoy, C.P., 117; 158
Page 540
Goldman, B., 426; 519

Goldman, J.C., 82; 102
See Tenore, K.R., 161
Goldsmith, H.L., 98; 102
See Karnis, A., 102
Gonor, J.J. See Barnes, H., 304, 324, 326; 385
González, M. See Jaramillo, E., 226
Gooch, J.L. See Snyder, T.P., 245, 246; 261
Gooding, R.M. See Ralston, S., 526
Gopalakrishna, V.V., 11, 16, 33, 58; 67
See Rao, R.R., 69
Gore, R.H., 137, 148; 155
Götting, K. See Gallardo, C.S., 250; 256
Goulet, D. See South, G.R., 529
Gouveia, A.D. See Shetye, S.R., 69
Grace, J.R. See Clift, R., 101
Graham, A. See Fretter, V., 236; 256
Graham, N.E. See Polovina, J.J., 525
Grahame, J., 235, 239, 240, 250, 251, 252, 289; 256, 295
See Crossland, S., 256
See Lowell, R.B., 258
Grand, S., 407, 410, 412, 422, 453; 519
Grandin, P., 487, 490, 491, 493, 494; 519
Grandperrin, R. See Brouard, F., 437, 439, 440; 514
See Kulbicki, M., 411, 439; 521
Grant, C., 213, 214; 226
Grassle, J.F., 246; 256
Grassle, J.P. See Grassle, J.F., 246; 256
Gray, W.N., 465, 466; 519
Green, R.H. See Singh, S.M., 244; 261
Greening, H.S., 136, 138, 139 155
Greenway, J., 213, 214; 226
Greenway, J.P. C., 210, 211, 213, 214; 226
Greenway, M., 120; 155
Gregg, M.C. See Brainerd, K.E., 92, 95; 101
See Shay, T.S., 95; 103
Gregory, S. See Li, Y.H., 91; 102
Grieve, C. See Kailola, P.J., 521
Grigg, R.W., 430; 519
See Atkinson, M.J., 430; 514
Groom, T.T., 370, 377 388
Gruet, Y., 195; 226
Gruvel, A., 304, 305, 306, 307, 308, 310, 312, 313, 318, 319, 320, 321, 322, 323, 326, 327, 334, 341, 343, 368, 373; 388
Gudmundsson, G.A. See Alerstam, T., 254, 293
GuérinGanivet, J. 307, 310, 382, 383; 388
Guille, A., 404; 519
Guillou, J., 194, 195, 197; 226
See Bayed, A., 223
Gulbrandsen, Ø., 433; 519
Gulland, J.A., 461; 519
Gundermann, N., 424; 519
Gunn, M.E. See Bleil, D.F., 281; 293
Gunnill, F.C., 148; 155
Gurney, R., 303; 388
Gwilliam, G.F., 322, 325, 328, 340, 380; 388
Gwyther, D., 471, 490, 495; 519
Gwyther, D.G. See Frusher, S.D., 519
Gwyther, J. See Munro, J.L., 483; 524
Haas, L.W. See Neckles, H.A., 158

Hacker, S.D., 149; 155
See Steneck, R.S., 160
Hacking, N. See McLachlan, A., 227
Haddon, M., 212, 214; 226
See Wear, R.G., 209, 213; 232
Haderlie, E.C., 167; 226
Hadlock, R.P., 272; 295
Haight, W., 440; 519
Haines, A., 445, 451; 519
Haines, A.K., 464, 467, 470, 492; 519
Haines, E.B., 282; 295
Hair, C., 490, 496; 520
Halapua, S., 407, 410, 412, 419, 420; 520
Hall, J.R., 280; 295
Hall, M.A., 187; 226
Hall, M.O., 117, 148; 155
Hallaher, E.E. See Gore, R.H., 155
Hallegraeff, G.M., 219; 226
Hallier, JP., 458; 520
Hamilton, L.S. See Wilson, A.M., 410; 531
Hamilton, P.V., 245, 280, 281, 282; 256, 295
See Dix, T.L., 282; 294
See Stirling, D., 280; 299
Hamm, D. See Craig, P., 516
Hamm, D.C., 410, 498; 520
Hanekom, N. See McLachlan, A., 227
HannafordEllis, C.J., 251; 256
Hansen, D.F. See GeertzHansen, O., 155
Happel, J., 85; 102
Harding, S.P. See Hawkins, S.J., 295
Hardwick, J.E. See Miller, D.J., 228
Hardy, J.T., 484; 520
Hardy, R. See Wendell, F., 232
Hardy, S.A. See Hardy, J.T., 484; 520
Harkantra, S.N., 205; 226
See Parulekar, A.H., 229
Harlin, M.M., 115, 116, 117, 132, 134; 155
Harnickell, E. See Walter, H., 70
Harrington, J. See Zimmerman, R.J., 162
Harrington, S.A. See Simpson, R.D., 249; 261
Harrison, P.G., 118; 155
Harrison, P.J. See Dortch, Q., 101
See Waite, A.M., 104
Hart, A., 251; 256
Hart, A.M., 423; 520
Hartnoll, R.G., 288; 295
See Hawkins, S.J., 120, 121, 122, 287; 155, 195
Hasse, J.J., 424; 520
Hassler, T.J. See Shaw, W.N., 165, 167, 168, 169, 170, 171; 231
Hastenrath, S., 5, 6, 24, 55; 67
Hastie, L.C., 489, 496; 520
Hatcher, A. See Williams D.M., 426; 531
Page 541
Hatcher, B.G., 400, 401; 520

Hawkes, W. See Kendrick, G.A., 117; 156
Hawkins, A.J.S. See Bayne, B.L., 121, 146; 153
Hawkins, S.J., 120, 121, 122, 265, 276, 287; 155, 295
See Davies, M.S., 294
See Hartnoll, R.G., 288; 295
See Imrie, D.W., 296
See Norton, T.A., 298
Hay, M.E., 141, 143; 155
See Holmlund, M.B., 156
Hay, T.K. See Boulding, E.G., 241; 255
Hayes, T.A. See Haight, W., 519
Hazlett, B. See Bach, C., 293
Heblekar, A.K. See Babu, M.T., 66
Heck, K.L., 139, 140, 141, 147, 148, 149; 155
See Coen, L.D., 154
See Orth, R.J., 159
See Valentine, J.F., 119, 145; 161
Hedgecock, D. See Mastro, E., 259
Hedgpeth, J.W. See Ricketts, E.F., 392
Heijs, F.M.L., 114, 115, 117, 118, 132, 133, 134, 150; 155, 156
See Brouns, J.J.W.M., 114, 133, 135; 153
Heller, J., 238; 256
Helm, N., 410, 415, 431, 498; 520
Helmuth, B., 249; 256
Henry, D.P., 306, 312, 313; 389
Henry, R.P. See McBride, C.J., 297
Hensley, R.A., 413, 415, 417; 520
Hermelink, P.M. See Jacobs, R.P.W.M., 156
Herndl, G.J. See Peduzzi, P., 146, 150; 159
HeronAllen, E., 303; 389
Herrera, C.B. See Searcy Bernal, R., 231
Herrington, W.C., 169; 226
Herrnkind, W.F. See Wilber, T., 278, 281, 282; 301
Hesp, P. See McLachlan, A., 227
Hesse, O. See Sornin, J.M., 160
Hewatt, W.G., 309; 389
Heymans, J.J., 201; 226
Hicks, G.R.F., 122; 156
Higgins, H.W. See Tranter, D.J., 161
Highsmith, R.C., 246, 249; 257
Hilborn, R., 188, 501, 502; 226, 520
Hilgard, G.H., 309, 346, 361, 362, 364, 366, 367, 368, 369, 370, 371; 389
Hill, A. See Black, R., 293
Hill, A.S. See Hawkins, S.J., 295
Hill, E.M. See Song, W.C., 393
Hill, L. See Wright, A., 399, 483, 484, 500, 502; 531
Hill, P.S. See Jumars, P.A., 102
Hillman, K., 111, 114, 115; 156
Hinds, V.T., 465, 466; 520
Hirschhorn, G., 173, 174; 226
Ho, Y.B. See Bryan, G.W., 294
Hodgson, A.N., 201; 226
Hoedt, F., 460; 520
Hoek, P.P.C., 304, 305, 306, 307, 308, 334, 373; 389
Hoeksma, B.W. See Breeman, A.M., 288; 293
Hoffman, D.L., 355, 356, 357, 358, 359, 360, 379; 389
Hogan, A.E. See Lewis, A.D., 471; 522
Hogue, E., 173; 226
Holberton, R. See Helmuth, B., 256
Holland, A., 476; 520
Holland, D.L., 353; 389
See Song, W.C., 393
Holland, K.N., 425; 520
Hollingsworth, C.E. See Lewis, J.B., 137, 138, 148; 157
Holmes, H.B. See Weymouth, F.W., 232
Holmes, R.W. See Eppley, R.W., 101
Holmlund, M.B., 147, 149; 156
Holt, S.J. See Beverton, R.J.H., 496, 503; 514
Holter, A.R., 329, 330; 389
Hooper, A., 399, 410, 431, 498; 520
Hootsmans, M.J., 118, 122, 126, 141, 143, 149; 156
Hopkinson, C.S. See Kemp, P.F., 296
See Schubauer, J.P., 283; 299
Horner, S.M. J., 117, 133; 156
Horsley, D.T. See Clarke, B., 255
Howard, G.K., 344; 389
Howard, N. See Sims, N., 483, 484; 528
Howard, R.K., 109, 115, 119, 120, 122, 123, 125, 127, 136, 138, 139, 141, 142, 146, 149; 156
See Klumpp, D.W., 157
See Robertson, A.I., 138, 139; 159
See Virnstein, R.W., 136, 137, 148; 161
Hruby, T. See Lewin, J., 227
Hubbard, D.M. See Dugan, J.E., 225
Hudson, R.J. M. See Morel, F.M.M., 103
Hughes, D.A., 205; 226
Hughes, J.M., 238; 257
Hughes, R.G., 120; 156
Hughes, R.N., 245, 247, 250, 251, 252; 257
See Currey, J.D., 242; 256
See Knight, A.J., 258
See Lawton, P., 272; 296
Hui, E., 307, 339, 342, 358, 379; 389
Huisman, J.M. See Kendrick, G.A., 156
Hull, S.L. See Grahame, J., 256
Hulo, J., 407, 453; 520
Humm, H.J., 117, 132, 133, 135; 156
See Ballantine, D.L., 115; 153
Hummerston, L.G. See Bryan, G.W., 294
Hunt, D.C. See Berry, A.J., 247; 255
Hunt, H.L. See Metaxas, A., 297
Hunt, J.R. See Logan, E.L., 72, 77; 102
Hunte, W., 249; 257
Hunter, J.T. See Stiven, A.E., 281; 299
Hunter, R.D., 265; 295
Hutchings, P.A., 135, 136; 156
See Collett, L.C., 154
Page 542
Hutchings, P.J. See Howard, R.K., 156

Hutchinson, G.E., 84; 102
Hutchinson, S. See Hawkins, S.J., 295
Huxham, M., 253, 291; 257, 296
Hviding, E., 410, 411; 520
Hylleberg, G. See Janekarn, V., 13, 16; 67
Hylleberg, J., 240, 265, 272; 257, 296
See Sundstrom, B., 70
Ianelli, J., 462; 520

See Gillett, R., 406, 453, 455, 457; 519
Ikehara, I., 443, 457, 503; 520
Ikehara, I.I. See Tsuda, R.T., 530
Imrie, D.W., 265; 296
Ingola, B.S. See Parulekar, A.H., 229
Innes, A.J., 329, 349; 389
Ino, T., 237; 257
Intes, A., 480, 485; 520
Ireland, M.P., 282; 296
Irlandi, E.A., 121; 156
Ishiyama, V. See Salgado, I., 192; 230
Iskakov, M.S. See Kutateladze, S.S., 102
Itano, D. See Buckley, R., 515
Itano, D.G., 434, 443, 444, 506; 520
Ivanov, J.A., 23; 67
Jablonski, D., 246; 257

Jackson, G.A., 83; 102
Jackson, J.B. C., 140; 156
Jacobs, R.P.W.M., 114, 117, 118, 119, 132, 133, 135, 146; 156
James, R. See Frid, C.L. J., 276, 277, 278; 295
Janekarn, V., 13, 16; 67
See Sundstrom, B., 70
Janke, K., 271, 272, 273, 274, 284, 292; 296
Janson, K., 236, 238, 239, 240, 241, 244, 245, 247, 248; 257
See Ward, R.D., 236, 246; 262
Jantarapagdee, P. See Yesaki, M., 11, 21; 70
Janzen, D.H. See Orians, G.H., 286; 298
Jaramillo, E., 191; 226
See Defeo, O., 224
See McLachlan, A., 163–232; 227
Jayaraman, R., 10; 67
See Anand, S.P., 67
See Rao, L.V.G., 11, 23, 25, 34, 35, 44, 50, 52, 57, 59; 69
Jeffery, D.J., 85; 102
Jeffries, C. See Ramage, C.S., 68
Jenkins, M.D. See Wells, S.M., 400, 401; 531
Jennings, S., 410, 411, 426, 427, 428, 429; 520
Jernakoff, P., 109–62, 141, 146, 267, 268, 274, 276; 156, 296
See Underwood, A.J., 269, 270, 274; 300
Jirka, G.H. See Brumley, B.H., 96; 101
Jobert, C. See Pouchet, G., 326; 392
Johannes, R.E., 407, 410, 413, 431, 453, 504, 508, 509; 520, 521
See Hatcher, B.G., 520
See Ruddle, K., 431; 527
Johannesson, B., 235, 239, 240, 244, 251; 257
See Johannesson, K., 235, 236; 258
Johannesson, K., 235, 236, 238, 239, 241, 244, 245, 246, 248, 249; 257, 258
See Alerstam, T., 254, 293
See Johannesson, B., 235, 239, 244, 251; 257
See Tatarenkov, A., 239; 261
Johansen, K. See Fyhn, H.J., 388
See Petersen, J.A., 392
Johns, S. See Hughes, R.G., 156
Johnson, A.S. See Trussell, G.C., 262
Johnson, C.R., 249, 252, 289, 290, 291; 258, 296
See Chapman, A.R. O., 288, 294
Johnson, C. See Parker, T., 298
Johnson, G.D., 435; 521
Johnson, M.S., 248; 258
Johnson, M.W. See Sverdrup, H.U., 70
Johnstone, I.M., 118; 156
Joleaud, A., 304, 318, 322; 389
Jones, G.P., 503; 521
Jones, H.D. See Davies, M.S., 294
Jones, J.S., 237; 258
Jorgensen, B.B. See SandJensen, K., 160
Joslyn, A.R. See Blackstone, N.W., 278; 293
Joubin, L., 307, 310, 382, 383; 389
Juch, P.J.W., 272; 296
Judy, M.H. See Dudley, D.L., 281; 294
Jumars, P.A., 71, 72, 73, 74, 81, 90, 95, 100; 102
See KarpBoss, L., 71–107
See Shimeta, J., 103
Kadolsky, D. See Bandel, K., 237; 254

Kailola, P.J., 404, 455, 466, 467, 478, 487; 521
Kalabushkin, B.A., 238; 258
Kamatie, M. See Preston, G.L., 526
Kami, H.T. See Ikehara, I., 520
See Tsuda, R.T., 530
Kamiya, H., 329; 390
Kamykowski, D., 76, 80, 81, 83; 102
Kan, T.T. See Matsuoka, T., 492; 523
Karakasch, N.I., 304; 390
Karnis, A., 98; 102
KarpBoss, L., 71–107
See Jumars, P.A., 102
Kartavtsev, Y.P., 247; 258
Katkansky, S.C. See Warner, R.W., 167; 232
Katnik, S.E., 410, 418, 419, 420, 421, 422; 521
Kaufmann, R., 368; 390
Kava, V. See Tu’avao, T., 530
Kawamoto, P.Y., 460; 521
Kazama, T.K., 460; 521
Kearney, R.E., 459; 521
See Ellway, C.P., 462; 518
See Gillett, R.D., 421; 519
See Kleiber, P., 421; 521
Keenan, C. See Brown, I.W., 515
Page 543
Keller, S.R., 84; 102

Kemp, P., 242; 258
Kemp, P.F., 280, 283; 296
Kendrick, G.A., 111, 115, 116, 117, 132, 134; 156
See Hutchings, P.A., 156
See Walker, D.I., 161
Kennelly, S.J. See Underwood, A.J., 265; 300
Kenworthy, W.J. See Fonseca, M.S., 111; 155
Kenyon, G. See Cook, L.M., 240; 255
Keyte, P. See Saenger, P., 206, 207; 229
Khokiattiwong, S., 59; 67
Kikuchi, T., 121; 157
Kilburn, R.N., 245; 258
Kilham, P. See Titman, D., 83; 104
Killingley, J.S. See Newman, W.A., 307; 391
Kimmerer, W.J. See Fancett, M.S., 155
Kimura, M., 243; 258
King, J.M. See Alldredge, A.L., 139; 153
King, M., 441, 452, 456, 457, 459, 460, 490, 493; 521
King, M.G., 489, 496; 521
King, P.A. See McGrath, D., 276; 297
King, R.J. to West, R.J., 161
Kiørboe, T., 71; 102
Kirchman, D.L., 118; 157
See Logan, E.L., 91; 102
Kirkman, H., 110, 111, 114, 115, 120, 122, 123, 145; 157
Kirkpatrick, G.J. See Kamykowski, D., 102
Kitalong, A., 410, 415, 423, 424, 425, 428; 521
Kitting, C.L., 116, 123, 126, 133, 139, 141; 157
See Morgan, M.D., 114, 122, 126, 135, 144; 158
Kleiber, P., 421; 521
Klepal, W., 322, 323, 324, 327, 334, 343, 368, 369, 373, 374, 375, 376, 377, 378; 390
See Barnes, H., 367, 368; 385
Klumpp, D.W., 119, 120, 122, 123, 128, 129, 145; 157
See Robertson, A.I., 120; 159
Kneib, R.T., 281; 296
Knight, A.J., 236, 245, 246; 258
See Ward, R.D., 262
Knox, R.B. See Ducker, S.C., 154
Knudson, K.K., 410; 521
Koch, A.L., 73; 102
Koch, G., 407; 521
Koehler, R., 318, 320, 321, 322, 323, 326, 333, 334, 353, 354; 390
Koepfler, E.T. See Neckles, H.A., 158
Kohlmeyer, J., 280, 283; 296
Kohnke, G., 408; 521
Kojis, B.L. See Quinn, N. 467, 470; 526
See Quinn, N.J., 404, 464, 466, 467, 470, 488, 490, 493; 526
Kolla, V., 58; 67
Konno, K. See Yoshino, T., 361; 394
Korson, L., 82; 102
Kostlan, E. See Smith, M.K., 440; 529
Kraeuter, J.N., 271; 296
Kramer, H., 82; 102
Krebs, C., 484; 521
Krishna Rao, P., 57; 67
Krohn, A., 321, 373; 390
Kronig, R., 82; 102
Krüger, P., 306, 308, 313, 319, 343, 366, 372; 390
Kuenzler, E.J. See Stiven, A.E., 282; 299
Kugele, M., 356, 357; 390
Kuhn, M. See McLachlan, A., 227
Kulbicki, M., 411, 426, 429, 439, 466, 467, 470; 521
See Baillon, N., 424; 514
See Hallier, J.P., 458; 520
See Wantiez, L., 471; 530
Kulczycki, G.R., 148; 157
Kuo, J., 110, 114; 157
See Cambridge, M.L., 114; 154
See Coles, R., 401; 516
Kusano, H., 147; 157
Kusano, T. See Kusano, H., 147; 157
Kutateladze, S.S., 85, 90; 102
Kyriakides, M.A. See Hawkins, S.J., 295
Laborda, A.J. See Mazé, R.A., 194, 197; 227

Laboute, P. See Guille, A., 519
Lacombe, D., 321; 390
La Fond, E.C., 10, 22, 24, 50, 51, 56, 57, 61; 67
See Ganapathi, P.N., 67
La Fond, K.G. See La Fond, E.C., 10, 24, 57; 67
Lagardère, F. See Ansell, A.D., 195, 197; 222
Lakshmana Rao, G.S. See Murty, V.S. N., 68
Lamb, P.J. See Hastenrath, S., 5, 6, 24, 55; 67
Landsberg, H.E., 4, 5; 67
Lang, S. See Wells, F.E., 161
LanganCranford, K.M., 236; 258
Langi, S. See Shomura, R.S., 528
Langi, S.A., See Langi, V.A., 433; 522
Langi, V.A., 433, 442; 522
Langston, W.J. See Bryan, G.W., 294
Lanyon, J., 119; 157
Lanzing, W.J.R. See Conacher, M.J., 154
Larkum, A.W.D., 111, 120; 157
See Conacher, M.J., 154
See Hillman, K., 156
See West, R.J., 161
Lassen, H.H. See Levinton, J., 238; 258
Lassuy, D., 172, 173, 174, 175, 176, 177, 178, 220; 227
Lastra, M., 199, 200; 227
Latu, S., 433, 439, 441, 503, 506; 522
Lauckner, G., 264; 296
Lavie, B., 244; 258
See Nevo, E., 298
See Noy, R., 259
La Violette, P.E., 11, 23, 26, 27, 38; 68
Lawrence, J.M., 120; 157
Page 544
Lawton, P., 272; 296

Layerle, C. See Defeo, O., 224
Layrisse, M. See Penchaszadeh, P.E., 229
Lazier, J.R. N., 71, 72, 82, 90, 92, 95; 102
See Mann, K.H., 71, 82; 103
Leal, G.L., 76, 78, 79, 85, 88, 98, 106; 102
Leary, D.F., 459, 460; 522
Leblic, I., 410; 522
LeClair, L.L., 174; 226
Ledua, E. See Adams, T.J. H., 512
Legeckis, R., 58; 68
Legendre, R. See GuerinGanivet, J. 307, 310, 382, 383; 388
Leighton, D., 237; 258
Lein, T.E., 272, 273, 279; 296
Leipper, D.F., 68
Leith, B.H. See Jones, J.S., 258
LeMoal, Y., 197; 227
See Guillou, J., 194, 195, 197; 226
Le Philippe, V. See Petit, D., 462; 525
Leqata, J. See Adams, T.J.H., 512
See Nash, W., 524
Leqata, J.L., 410, 411; 522
Lera, M.E. See Penchaszadeh, P.E., 229
Lessard, E.J. See Shimeta, J., 103
Lessios, H.A., 243; 258
Lethbridge, R. See Nielsen, J., 119, 121, 128, 138, 139, 141, 142, 146; 159
Lethbridge, R.C., 117; 157
See Borowitzka, M.A., 111, 114, 115, 116, 117, 118, 132, 133, 134, 135, 141; 153
Levinton, J., 238; 258
Levinton, J.S. See Lopez, G.R., 283; 296
Lewin, J., 173; 227
Lewis, A.D., 399, 433, 446, 449, 450, 451, 452, 458, 460, 466, 470, 471, 472, 478, 479, 480, 481,482; 522
See Adams, T.J. H., 512
See Chapman, L.B., 449; 515
See Dalzell, P., 456, 458, 462; 517
Lewis, C.A., 337, 341, 342, 344, 346, 358, 359, 360, 361, 362, 363, 364, 365, 366, 368, 369, 370, 371, 372, 373, 376, 377; 390
See Chaffee, J., 353, 355; 386
Lewis, F.G., 136, 137, 148; 157
See See Stoner, A.W., 137, 148, 149; 160
See Virnstein, R.W., 161
Lewis, J.B., 137, 138, 148; 157
Lewontin, R.C., 243; 258
Li, Y.H., 91; 102
Liceaga, M. See Defeo, O., 224
Lichatowich, T. See Gundermann, N., 519
Lighthill, J., 86; 102
Ligouri, V.R. See Lacombe, D., 321; 390
Limpus, C.J. See Lanyon, J., 157
Lindberg, D.R. See Chaffee, C., 317; 386
Lindholm, R. See Frusher, S.D., 519
Lindsay, S., 452; 522
Lindsay, S.R.. See Dalzell, P., 517
Link, T., 174, 175; 227
Lippman, H. See Landsberg, H.E., 67
Little, C., 267; 296
Littler, D.S. See Littler, M.M., 116, 121, 285; 158, 296
Littler, M.M., 116, 121, 285; 158, 296
See Seapy, R.R., 308; 392
Liu, L.L., 246; 258
Livingston, R.J. See Dugan, P.J., 138, 139, 140; 154
See Greening, H.S., 136, 138, 139; 155
Llabador, F., 308, 382; 390
Lochmann, S. See Jackson, G.A., 83; 102
Lock, J.M., 410, 411, 415, 418, 419, 420, 421, 428, 430, 446, 449; 522
Logan, E.L., 72, 77, 91; 102
See Confer, D.R., 72; 101
Lokani, P., 477, 482; 522
See Adams, T.J. H., 512
See Chapau, M.R., 410, 421, 428; 515
See Preston, G.L., 477; 526
Lopez, G.R., 283; 296
López, A., 192; 227
Lorenti, M. See Gambi, M.C., 155
Losada, F. See Penchaszadeh, P.E., 180; 229
Lou, D.C., 423, 424, 425; 522
Loubens, G., 410, 411, 412, 423, 424, 425, 502; 522, 523
Lowe, C.G. See Holland, K.N., 520
Lowell, R.B., 240; 258
See Mercurio, K.S., 259
Lubchenco, J., 121, 266, 271, 272, 273, 274, 275, 276, 285, 286, 287, 288; 158, 296, 297
Lucas, A., 195, 196; 227
Lucas, J.S., 483; 523
See Copland, J.W., 483; 516
See Robertson, A.I., 143; 159
Lucas, M.I. See Matthews, S., 227
Ludwig, G.M. See Ralston, S., 526
Lueck, R. See Osborn, T.R., 95; 103
See Yamazaki, H., 95; 104
Lukatelich, R.J., 150; 158
Lumley, J.L. See Tennekes, H., 92, 93; 104
Luther, M.E. See Potemra, J.T., 68
Lutz, R.A. See Jablonski, D., 246; 257
Lymbery, A. See Black, R., 293
Lyngby, J.E. See Brix, H., 117; 153
Lyonnet, A. See Defeo, O., 224
MacArthur, G.J. See Smith, P.J., 231

MacArthur, R.H., 249; 258
MacDonald, C.D., 491, 493, 494; 523
MacFarlane, J.W., 494; 523
See Bell, R.S., 514
See Johannes, R.E., 410; 521
Page 545
See Moore, R., 494; 524

MacIntyre, R.J., 358; 391
MacLean, A. See Shelford, V.E., 393
Maddock, J.R., 79; 102
Magoon, C.D. See Tegelberg, H.C., 173, 174, 178; 231
Mahmoud, M.F., 318, 319, 333, 354, 355; 391
Majkowski, J. See Shomura, R.S., 528
Malaika, J. See McNown, J.S., 84; 103
Mallet, A.L., 248; 258
Malone, T.C., 97; 102
Mane, U.H. See Talikhedkar, P.M., 231
Manley, N.L. See Norton, T.A., 298
Manly, R. See Fretter, V., 247, 289; 256, 295
Mann, K.H., 71, 82, 97; 102, 103
See Johnson, C.R., 249, 252, 289, 290, 291; 258, 296
See Lazier, J.R. N., 71, 72, 82, 90, 92, 95; 102
See Robertson, A.I., 109, 119, 123, 125, 139, 140, 141, 142, 147, 150, 251, 280, 283; 159, 260, 299
See Schneider, F.I., 141, 148; 160
Mansour, R.A. See Beherens Yamada, S., 277, 279; 293
Marais, J.F. K. See du Preez, H.H., 225
Marchionni, V. See Badino, G., 195; 222
Margalef, R., 97; 103
Markham, J. See Denton, A.B., 294
Marples, T.G., 282; 297
Marsh, C.P., 379; 391
Marsh, G.A., 136, 137, 138, 139, 140; 158
Marsh, H. See Lanyon, J., 157
Marshall, N. See Stevenson, D.K., 428; 529
Martel, A., 426, 429, 290; 259, 297
Martinez, R. See Santelices, B., 147; 160
Maruyama, T. See Nei, M., 259
Masello, A., 187; 227
See Defeo, O., 224
Masini, R.J., 118; 158
Maslennikov, V.V., 17, 19, 21, 50, 57; 68
Masliyah, J.H., 79, 85; 103
Mason, G.S. See Karnis, A., 102
Mason, S.G. See Forgacs, O.L., 95; 101
See Goldsmith, H.L., 98; 102
See Van de Yen, T.G.M., 95; 104
Massé, H., 195; 227
See Bodoy, A., 195, 197; 223
Mastro, E., 236; 259
Mataia, N. See Nash, W., 524
MatengaSmith, T., 498; 523
Mather, P.B. See Hughes, J.M., 238; 257
Mathew, B. See Viswambharan, N.K., 24; 70
Mathews, E., 407, 410, 413, 422, 472, 473, 474, 475, 479, 481, 492; 523
Mathieson, A.C. See Fralick, R.A. F., 295
Matsuoka, T., 492; 523
Matteo, M.B. See Vuilleumier, F., 236; 262
Matthews, S., 200, 201; 227
May, V., 111, 135; 158
Mazé, R.A., 194, 197; 227
Mazellier, P. See Bagnis, R., 514
Mazzella, L., 109, 110, 114, 117, 118, 119, 120, 123, 128, 133, 134, 135, 141, 142, 146, 147, 149; 158
See Casola, E., 154
See Cinelli, F., 154
See Kirchman, D.L., 157
McBride, C.J., 281; 297
McCollum, S.A. See Kamykowski, D., 81, 83; 102
McComb, A.J., 115, 132; 158
See Cambridge, M.L., 118, 150; 154
See Kendrick, G.A., 156
See Kuo, J., 110, 114; 157
See Larkum, A.W. D., 157
See Lukatelich, R.J., 158
See Masini, R.J., 158
See Silberstein, K., 160
See Walker, D.I., 161
See Shepherd, S.A., 160
McConnell, O.J. See Geiselman, J.A., 279, 286; 295
McCook, L.J., 275, 288, 289, 292; 297
McCoy, M., 418, 438; 523
McCrohan, C.R. See Imrie, D.W., 296
McFadyen, K.E. See Underwood, A.J., 247, 250, 267, 276; 262, 300
McGrath, D., 276; 297
McGraw, K.A. See Schink, T.D., 230
McGregor, D.B., 327; 391
McIntosh, C.R. See Rapson, A.M., 405; 527
McKillup, S.C., 268; 297
McKinnon, S., 209, 210, 211, 212; 227
McLachlan, A., 163–232, 167, 172, 186, 199, 200, 201, 202, 205, 206, 207, 209, 213, 215, 217; 227
See Ansell, A.D., 201; 222
See Cockcroft, A.C., 201; 223
See Donn Jr, T.E., 225
See du Preez, H.H., 225
See Heymans, J.J., 201; 226
See Prosch, R., 201; 230
McLaughlin, J.B., 98, 99; 103
McLean, R.B., 278, 282; 297
McMillan, C. See Zapata, O., 118; 162
McMillin, H.C., 172, 173, 174; 228
See Weymouth, F.W., 173; 232
McNee, A. See Kailola, P.J., 521
McNown, J.S., 84; 103
McPherson, G. See Brown, I.W., 515
McPherson, G.R., 423, 424, 450, 502; 523
McQuaid, C.D., 233–62, 263–302, 234, 239, 240, 245, 247, 248, 252, 253, 254, 264, 268, 277, 280, 281, 283, 284, 291, 292, 259, 297
McRoy, C.P., 117; 158
McShane, P. See Black, R., 293
Page 546
McVey, J.P. See Hasse, J.J., 520

Mead, P., 439; 523
Mead, P.D. See Preston, G.L., 526
Meekan, M., 249; 259
Mees, C.C., 410, 418, 428, 438, 440, 442, 453; 523
Meese, R.J., 357, 358, 379, 380; 391
Menge, B.A., 272, 274, 276; 297
See Lubchenco, J., 271, 272; 297
Menou, J.L. See Guille, A., 519
Mercurio, K.S., 238; 259
Mestayer, M.K., 211, 212; 228
Metaxas, A., 288; 297
Metz, N.J., 306; 391
Meyer, G.R. See O’Gower, A.K., 269; 298
Michael, G.S. See Shetye, S.R., 69
Michael, K.P. See Cranfield, H.J., 223; 224
See Smith, P.J., 231
Mierle, G., 75; 103
Mileikovsky, S.A., 249, 250; 259
Milicich, M.J. See Meekan, M., 259
Milkman, R., 237; 259
Mill, P. See Grahame, J., 240; 256
Mill, P.J. See Crossland, S., 256
See Grahame, J., 240, 252; 256
See Lowell, R.B., 255
Millar, R.B., 209, 210; 228
Miller, D. See Coates, D., 516
Miller, D.J., 167, 170; 227
Miller, F.R. See Ramage, C.S., 68
Miller, G.H. See Salvigsen, O., 299
Miller, M. See Leary, D.F., 522
Miller, S.L., 274; 297
Millero, J.F. See Korson, L., 102
Milton, D.A., 456, 460; 523
See Blaber, S.J. M., 404, 466; 514
See Rawlinson, N.J. F., 527
Mitchell, B.D. See Barnes, H., 385
Mitchell, J.G., 84; 103
Mitchell, R. See Kirchman, D.L., 157
Mitchum, G.T. See Polovina, J.J., 493; 525
Mix, M.C., 334; 391
See Abbott, D.T., 381
Miyamoto, G.T. See Ralston, S., 440; 526
Mizrahi, L. See NeubergerCywiak, L., 228
Mobiha, A., 423, 451; 523
Moce, K. See Cooke, A., 432; 516
Moffitt, R.B., 443; 523
Mojumdar, P., 11; 68
Molares, J., 362, 366, 372, 373, 383; 391
Molinari, R.L., 59, 62; 68
Moltschaniwskyj, N.A. See Lou, D.C., 423, 424; 522
Monin, A.S., 92; 103
Mook, D., 124; 158
Moore, D.G. See Kolla, V., 67
Moore, H.B., 267, 272, 276; 297
Moore, K.A. See Orth, R.J., 115, 132; 159
Moore, L. See Cambridge, M.L., 154
Moore, R., 471, 494; 523, 524
See Bell, R.S., 514
See MacFarlane, J.W., 494; 523
See Reynolds, L.F., 471; 527
MoralesNin, B., 424, 502; 524
Moran, M.J., 267; 297
See Fairweather, P.G., 295
See Underwood, A.J., 300
Morel, F.M. M., 75; 103
Moreno, V. See Olivier, S., 229
Morgan, G.R. See Pauly, D., 501; 525
Morgan, M.D., 114, 120, 122, 124, 126, 135, 144; 158
See Kitting, C.L., 157
Morize, E., 410, 412, 425, 453; 524
See Caillart, B., 419; 515
Morris, R.J., 350, 351, 352; 391
Morris, S.R. See Beardmore, J.A., 235, 244; 254
MouTham, G. See Kulbicki, M., 521
Mouëza, M., 194, 195, 196, 197; 228
See Ando, T., 222
See Degiovanni, C., 195, 197; 224
See Frenkiel, L., 196; 226
See Pichon, Y., 229
Moynihan, E. See Wilkins, N.P., 262
Moyse, J., 236, 373; 259, 391
See Hui, E., 358; 389
Mueller, G.J. See Metz, N.J., 306; 391
MuellerDombois, D. See Walter, H., 70
Muller, R.G., 458, 459, 460, 461, 462, 503; 524
Munk, W.H., 71, 82, 84; 103
Munn, E.A., 367; 391
See Klepal, W., 390
Munro, D. See Nash, W., 524
Munro, I.S. R., 450; 524
Munro, J.L., 399, 410, 418, 422, 427, 430, 483, 484, 493, 494, 498; 524
See Pauly, D., 168, 169; 229
See Pearson, R.G., 483; 525
Murawski, S.A., 219; 228
Murphy, G.I. See Leary, D.F., 459, 460; 522
Murthy, C.B. See Anand, S.P., 66
Murty, C.S., 10, 19; 68
See Antony, M.K., 66
See Babu, M.T., 66
See Murty, V.S. N., 68
See Premchand, K., 68
See Rao, K.H., 69
See Varadachari, V.V.R., 70
Murty, S.V. See Ganapathi, P.N., 10; 67
Murty, V.S.N., 16, 23, 24, 33, 38, 43, 44, 51, 52, 60, 64; 68
See Gopalkrishna, V.V., 67
See Rao, D.P., 64; 69
Page 547

See Varkey, M.J., 1–70
Muyard, J., 408; 524
Mwaidseje, B., 204; 228
Myers, R.F., 399, 404, 410, 413; 524
Nagabhushanam, R., 205, 206; 228

See Talikhedkar, P.M., 231
Nagle, J.S., 136, 137, 138, 140; 158
Nair, S.A. See Otero, J.F., 229
Nakamura, E., 459; 524
Nakoryakov, V.E. See Kutateladze, S.S., 102
Naldrett, M.J., 321; 391
Nampoothiri, G. See Shetye, S.R., 69
Naqvi, S.W.A. See De Souza, S.N., 67
Narchi, W., 180, 181, 187, 188; 228
Naseli, N. See Preston, G.L., 526
Nash, W., 479, 484, 498; 524
Nath, G. See Dalzell, P., 517
Navaz, J.M., 307; 391
Nayar, K.N., 205, 206; 228
Naylor, R., 238, 240, 245; 259
Nebert, M. See McRoy, C.P., 258
Neckles, H.A., 109, 110, 117, 119, 123, 129, 130, 141, 149, 150; 158
See Wetzel, R.L., 119; 161
Nei, M., 243, 244; 259
Nejman, V.G., 56; 68
Nelson, G. See Whitehead, P.J.P., 531
Nelson, W.G., 137, 138, 139, 140, 141, 142, 149; 158
See Kulczycki, G.R., 157
See Virnstein, R.W., 161
NeubergerCywiak, L., 197; 228
Neushul, M. See Foster, M., 388
Neverauskas, V. See Shepherd, S.A. 160
Nevo, E., 264; 298
See Lavie, B., 244; 258
See Noy, R., 259
Newbury, S.F. See Atkinson, W.D., 250; 254
Newell, G.E., 272; 298
See Smith, J.E., 245; 261
Newell, R.C., 267; 298
Newell, S.Y. See Barlocher, F., 283; 293
See Kemp, P.F., 296
Newkirk, G.F., 236, 240; 259
Newman, R.A. See Rickard, N.A., 178; 230
Newman, W.A., 304, 305, 306, 307, 311, 312, 359, 361, 364, 381, 382; 391, 392
Nguyen, F. See Carlot, A.H., 434; 515
See Schaan, O., 528
NguyenKhoa, S., 411; 524
Nichols, P.D. See Klumpp, D.W., 119, 120; 157
Nichols, P.V., 410, 458, 462, 488, 492, 498; 524
See Blaber, S.J.M., 514
See Evans, D., 458, 460; 518
See Leqata, J.L., 522
Nickerson, R.B., 172, 174, 175; 228
Nicotri, M.E., 122, 123, 136, 142, 265; 158, 298
Nielsen, C. See Ockelmann, K.W., 247, 289, 290; 259, 298
Nielsen, J., 119, 121, 128, 137, 138, 139, 140, 141, 142, 146; 158, 159
See Jernakoff, P., 109–62
Nielson, J.D. See Campana, S.E., 502; 515
Nienhuis, P.H., 122; 159
Nieuwolt, S., 6; 68
NilssonCantell, C.A., 305, 306, 308; 392
NilssonCantell, C.A., 304, 306, 308, 312, 313, 334, 340, 343; 392
Nishikawa, S., 369; 392
Nobré, A., 308; 392
Norton, T.A., 143, 265, 271, 273, 279, 289, 290; 159, 298
See Hawkins, S.J., 295
See Watson, D.C., 273, 274, 276, 284, 285, 286, 287; 301
Noten, T.M.P.A. See Jacobs, R.P.W.M., 114, 118, 133, 135; 156
Novak, R., 133; 159
Noy, R., 244; 259
See Nevo, E., 298
Nussbaum, M., 318, 321, 322, 323, 324, 327, 334, 339, 343, 370, 371, 373, 374, 375; 392
Nybakken, J., 165, 166, 168; 229
Oakey, N.S., 92; 103

O’Bryan, P.G. See Tsuda, R.T., 530
O’Brien, J. See Mcpherson, G.R., 523
O’Brien, J.J. See Potemra, J.T., 68
See Yu, L., 70
Ockelmann, K.W., 247, 289, 290; 259, 298
Odum, E.P. See Odum, H.T., 426; 524
Odum, H.T., 426; 524
Ó Foighil, D., 249; 259
Ogata, K. See Kamiya, L., 329; 390
Ogden, J.C., 120; 159
See Vadas, R.L., 161
O’Gower, A.K., 269; 298
Ohlhorst, S.L., 139; 159
Ohta, T., 243, 247; 259
Oiterong, E. See Mathews, E., 407, 410, 413, 422, 472, 473, 474, 475, 479, 481, 492; 523
Olesen, B., 111; 159
Olive, P.J.W., 251, 252; 259
Olivier, S., 186, 187, 188, 189, 190; 229
See Penchaszadeh, P.E., 188; 229
Olsen, D. See McKinnon, S., 209, 210, 211, 212; 227
See Millar, R.B., 209, 210; 228
See Molinari, R.L., 68
O’Neill, M.E. See Dorrepaal, J.M., 85; 101
Opnai, L.J., 464, 465, 470, 494; 524, 525
See Evans, C., 492, 495; 518
See Polovina, J.J., 495; 525
O’Regan, D. See Wilkins, N.P., 247; 262
Page 548
Orians, G.H., 286; 298

Orth, F.L., 178; 229
Orth, R.J., 109, 110, 115, 117, 118, 119, 132, 141, 142, 147, 149, 152; 159
See Heck, K.L., 141, 147, 148; 155
See Neckles, H.A., 158
See Van Montfrans, J., 161
Ortiz, E. See Defeo, O., 224
Osborn, T.R., 95; 103
Otero, J. See Fernandez, J., 225
Otero, J.F., 195, 196, 197, 198; 229
Ott, J., 114, 115, 120, 132; 159
See Mazzella, L., 114, 146; 158
Oviatt, C.A. See Doering, P.H., 146; 154
Padilla, D.K., 121, 285; 159, 298

Paffen, K.H. See Landsberg, H.E., 67
Page, F.H. See Thomas, M.L.H., 289, 290; 300
Page, H.M., 306, 345, 346, 347, 349, 358, 359, 360, 361, 362, 363, 364, 365, 366; 392
Paine, R.T., 308, 359, 380; 392
Palmer, A.R., 241, 249, 253, 379; 260, 392
See Appleton, R.D., 241, 242, 253; 254
See Mercurio, K.S., 259
Parada, E. See Peredo, S., 229
Parker, P.L. See Goering, J.J., 118; 155
Parker, T., 288; 298
Parkin, D.T. See Clarke, B., 255
Parkinson, B.J., 478; 525
Parkinson, J.S., 79; 103
Parrish, J.D. See Haight, W., 519
Parulekar, A.H. 204; 229
See Harkantra, S.N., 205; 226
Pasciak, W.J., 75, 91; 105; 103
Pascual, C. See Molares, J., 391
Passfield, K.D., 487; 525
Patiale, H., 410, 418, 498; 525
See Dalzell, P., 517
Paulo, T.A. See Zoutendyk, D., 498; 531
Pauls, S.M., 184; 229
Pauly, D., 168, 169, 432, 501, 505; 229, 525
Pauly, D. See Gayanilo, F.C., 226
Pearse, J.S. See LanganCranford, K.M., 236; 258
Pearson, R.G., 483; 525
Peckol, P. See Yates, J.L., 286; 302
Pednekar, S.M. See Gopalkrishna, V.V., 67
Peduzzi, P., 121, 127, 141, 142, 144, 146, 150; 159
Pelseneer, P., 196; 229
Penchaszadeh, P. See Olivier, S., 186, 190; 229
Penchaszadeh, P.E., 180, 181, 184, 188; 229
See Diaz, M.C., 225
See Pauls, S.M., 229
Penhale, P.A., 117, 119, 132, 135, 149; 159
Penn, J.S. See Hamilton, P.V., 295
Peperzak, L. See Van Ierland, E.T., 81; 104
Peredo, S., 192; 229
Pérès, J.M. See Kikuchi, T., 121; 157
Perez, O. See Carriquiriborde, L., 223
Perrine, D., 488, 492, 495; 525
Peters, R.S., 280; 298
Petersen, J.A., 329, 331, 348, 349; 392
See Fyhn, H.J., 388
Peterson, C.H., 218, 249; 229, 260
See Holmlund, M.B., 156
See Irlandi, E.A., 121; 156
Peterson, J.D. See Holland, K.N., 520
Petit, D., 462; 525
Petraitis, P.S., 245, 270, 272, 273, 274, 275, 276, 277; 260, 298
Petriconi, V., 322, 343; 392
Pettitt, C., 238, 266; 260, 298
Pfister, C.R. See Hay, M.E., 155
Phelps, S.R. See LeClair, L.L., 174; 227
See Shaklee, J.B., 528
Phillips, B.F. See Bell, R.S., 514
Pichon, Y., 197; 229
Picken, G.B., 249; 260
Pike, A. See Huxham, M., 257, 296
Pilkington, M.C., 252; 260
Pilsbry, H.A., 304, 305, 306, 307, 313; 392
Pino, M. See Jaramillo, E., 226
Pita, E., 474; 525
Pitcher, R., 489, 490, 493, 494; 525
Plante Cuny, M.R., 197; 229
Poli, G., 195; 229
Pollard, D.A., 119; 159
See Bell, J.D., 120, 147; 153
See Klumpp, D.W., 157
Pollard, R.T., 53; 68
Polovina, J.J., 403, 430, 432, 440, 441, 442, 491, 493, 495, 500, 502; 525
See Grigg, R.W., 519
See Langi, V.A., 522
See Ralston, S., 432, 441; 526
Polunin, N.V.C., 430, 500; 525
See Dalzell, P., 395–531
See Jennings, S., 410, 411, 426, 427, 428, 429; 520
Ponwith, B. See Craig, P., 516
Poore, G.C.B., 147; 159
Poornachandra Rao, C., 56; 68
Popper, D.M. See Gundermann, N., 519
Potemra, J.T., 60, 61; 68
Pouchet, G., 326; 392
Poupin, J., 489, 496; 525
Prasanna Kumar, S. See Babu, M.T., 66
See Somayajulu, Y.K., 70
Premchand, K., 52; 68
Prenant, M., 307, 310, 318, 354; 392
See FischerPiette, E., 307, 308, 310, 382; 387
Prescott, J., 486, 487, 490, 491, 493, 494, 498; 525, 526
Press, W.H., 107; 103
Preston, G.L., 404, 413, 445, 473, 474, 475, 477, 478, 479, 486; 526
Page 549
See Dalzell, P., 406, 417, 418, 432, 435, 438, 439, 440, 443; 517
See Fusimalohi, T., 439; 519
See Sims, N., 528
Price, N.M. See Morel, F.M. M., 103
Prieto, A.S., 180, 181, 182; 229, 230
Pring, C.K. See Lewis, A.D., 446, 470, 472; 522
Prosch, R., 201; 229
Purcell, E.M., 72, 84, 89, 90, 95, 99; 103
See Berg, H.C., 71, 72, 78, 82, 83, 107; 101
Pye, V.I. See Newell, R.C., 298
Pyle, R.L., 422, 431; 526
Quach, M.M.C. See Hamm, D.C, 520

Quayle, D.B., 174; 229
See Bourne, N., 172, 174; 223
Quinn, G.P., 269, 270, 277; 298
See Braley, H., 293
See Ward, S., 268; 301
Quinn, N., 407, 465; 526
Quinn, N.J., 404, 464, 466, 467, 470, 488, 490, 493; 526
Quintana, R., See Molares, J., 391
Radhakrishna, K., 10; 68

Radtke, R.L. See Dee, A.J., 425; 517
Raffaelli, D., 235, 240, 264; 260, 298
See Huxham, M., 257, 296
Raftery, R.E., 280; 298
Raimbault, R. See Comps, M., 196; 223
Rainer, S. See Poore, G.C.B., 147; 159
Ralston, S., 423, 424, 425, 432, 440, 441, 496, 502; 526
See MoralesNin, B., 424, 502; 524
See Polovina, J.J., 440, 441, 500, 502; 525
Rama Raju, V.S. See Rao, N.T.V., 69
Ramage, C.S., 5, 6, 8; 68
Ramana Murty, T.V. See Somayajulu, Y.K., 70
Ramasastry, A.A., 10; 68
See Balaramamurty, C.B., 56; 66
Ramesh Babu, V., 17, 18, 23, 52, 57; 68
Rammer, A. See Rickard, N.A., 230
Ramohia, P. See Adams, T.J.H., 512
Randall, J.E., 423; 526
See KuMcki, M., 521
Rangeley, R.W., 272, 287; 298
Rao, D.P., 11, 23, 27, 28, 30, 39, 40, 48, 49; 69
See Babu, M.T., 66
See Murty, V.S.N., 68
See Rao, N.T.V., 69
Rao, K.H., 57, 69
See Antony, M.K., 66
Rao, K.L., 7; 69
Rao, K.N., 6, 8; 69
Rao, K.S., 205; 230
Rao, L.V.G. 11, 23, 25, 34, 35, 44, 50, 52, 57, 59; 69
See Sarma, M.S.S., 58; 69
Rao, N.T.V., 11; 69
Rao, P.D. See Rao, N.T.V., 69
Rao, R.R., 23; 69
Rao, T.S.S. See Ganapathi, P.N., 10; 67
Rao, Y.P., 5, 6; 69
Rapson, A.M., 209, 211, 405, 454; 230, 526, 527
Rasmussen, D.I. See Shelford, V.E., 393
Rastelli, M.L. See Sacchi, C.F., 235; 261
Rata, T., 457; 527
Raven, J.A., 84; 103
Rawlings, P. See Jones, J.S., 258
Rawlinson, N.J.F., 407, 410, 411, 427, 458, 459, 460, 466, 470, 472, 479, 481; 527
See Blaber, S.J.M., 514
See Milton, D.A., 523
See Nichols, P.V., 458, 462, 498; 524
Reddy, C.V.G. See De Souza, S.N., 67
See Sankaranarayan, V.N., 27, 36; 69
See Varadachari, V.V.R., 70
Reddy, G.V. See Antony, M.K., 66
See Rao, K.H., 69
Redfearn, P., 208, 209, 210, 211, 212, 213, 214; 230
Reed, R.E. See Kamykowski, D., 102
Reese, E.S. See Barnes, H., 304, 306, 308, 321, 328, 329, 335, 337, 338, 339, 340, 341, 342, 345, 355, 359, 361, 363, 365, 368, 380; 385
Reichel, E. See Baumgartner, A., 6, 7, 8; 66
Reichelt, J.L., 118; 159
Reichelt, R.E. See Kailola, P.J., 521
Reid, D.G., 234, 235, 236, 238, 239, 242, 248, 249, 250, 252, 253, 264, 265, 267, 269; 260, 298, 299
Reimchen, T.E., 237, 238, 245, 250, 253, 284, 287; 260, 299
Reise, K. See Wilhelmsen, U., 271, 273; 301
Renault, L. See Mouëza, M., 195; 228
Reverdin, G. See Molinari, R.L., 68
Revsbach, N.P. See SandJensen, K., 160
Rey, M. See Defeo, O., 224
Reynolds, L.F., 471; 527
See Moore, R., 471; 524
Rice, J.D. See Trocine, R.P., 161
Rice, L., 306; 392
Rice, L.A. See Shelford, V.E., 393
Rich, W.H. See Weymouth, F.W., 232
Richard, G., 479, 481, 482, 483, 484; 527
Richards, A., 412, 431, 473, 497; 527
See Adams, T.J.H., 512
See Sundberg, P., 438, 529
Richards, A.H., 418, 438, 439, 440, 490, 498; 527
See Wright, A., 410, 418, 419, 421, 428, 446, 449; 531
Richards, K.S. See Drew, K.M., 378; 387
Richardson, J.R., 207, 213, 215; 230
Richardson, K. See Raven, J.A., 84; 103
Page 550
Richmond, R.H., 476; 527

Rickard, N.A., 173, 178; 230
Ricker, W.E., 427, 429, 484, 503; 527
Ricketts, E.F., 306, 308, 317; 392
Riebesell, U., 75; 103
Riera, A., 184; 230
Riley, G.A. See Munk, W.H., 71, 82, 84; 103
Rincón, A. See Penchaszadeh, P.E., 181, 229
Rincón, A.B., 180, 181; 230
Rios, E., 182; 230
Rittschof, D., 282; 299
See Bach, C., 293
Robblee, M.B. See Zieman, J.C., 162
Roberts, A.M., 71, 72, 75, 82; 103
Roberts, C. See Polunin, N.V.C., 500; 525
Roberts, D.J. See Hughes, R.N., 247, 250, 251, 252; 257
Robertson, A.I., 109, 119, 120, 123, 125, 138, 139, 140, 141, 142, 143, 147, 149, 150, 251, 280, 283; 159, 260, 299
See Edgar, G.J., 136, 137, 139, 141, 147, 148, 149; 155
See Hatcher, B.G., 520
Robertson, C.H., 489; 527
Robinson, A. See Breese, W.P., 172; 223
Robles, A. See Arntz, W.E., 222
Rochford, D.J., 22, 51, 52; 69
Rodríguez, G., 180; 230
Rodríguez, S. See Molares, J., 391
Rohsenow, W.M., 77; 103
RolanAlvarez, E. See Johanneson, K., 258
Rosa, F. See Adams, T.J.H., 512
Rose, R.A. See Wells, F.E., 161
Rosenberg, A.A., 501; 527
Rosewater, J., 234, 249; 260
Ross, A. See Newman, W.A., 304; 391, 392
Roux, A., See Bastida, R., 223
Rowe, F. See Tranter, D.J., 161
Roy, J.M. See Parulekar, J.F., 229
Ruckelshaus, M.H. See Williams, S.L., 110, 122, 130, 141, 149, 151; 161
Ruddle, K., 431, 463; 527
Ruiz, C.A., 180; 230
Rudolph, S.G., See Trussell, G.C., 262
Russ, G., 427, 504; 527
See Brown, I.W., 515
See Ferreira, B.P., 425; 518
Russ, G.R. See Hart, A.M., 423; 520
RussellHunter, W.D. See Hunter, R.D., 265; 295
Russo, A.R., 148; 159
Russo, G.F., 139; 159
See Gambi, M.C., 155
See Mazzella, L., 123, 128, 141, 142, 146; 158
Ryan, N.R. See Quinn, G.P., 269, 270, 277; 298
Saavedra Rosas, A. See Searcy Bernal, R., 169; 231

Sacchi, C.F., 235; 238
Saenger, P., 206, 207; 230
Saffman, P.G., 98, 103
Sakamoto, R.K. See Amesbury, S.S., 512
Seelkehara, I., 520
SalasCasanova, C., 195; 230
Sale, P.F., 422, 500, 502; 527
Salemaa, H., 123, 144; 160
Salgado, I., 192; 230
Salini, J. See Shaklee, J.B., 528
SalitaEspinosa, J.T. See Klumpp, D.W., 157
Salvat, B., 195; 230
Salvigsen, O., 271; 299
Samoilys, M., 426; 527
See Brown, I.W., 515
Sanchez, J. See Defoe, O., 224
SandJensen, K., 118, 132, 133, 135, 150, 280; 160, 299
See GeertzHansen, O., 155
See Olesen, B., 111; 159
Sankaranarayan, V.N., 27, 36; 69
Santanam, K. See Murty, V.S. N., 68
Santelices, B., 141, 146, 147, 288; 160, 299
See Buschmann, A.H., 146; 154
Sarma, M.S.S., 58; 69
Sarma, Y.V.B. See Murty, V.S.N., 68
Sarsam, V.H. See Cook, L.M., 255
Sasabule, J.A., 410, 411; 528
Sasamal, S.K., 38; 69
Sastry, J.S., 38, 43, 51, 52; 69
See Gopalkrishna, V.V., 11, 16, 58; 67
See La Fond, E.C., 22; 67
See Premchand, K., 68
See Ramesh Babu, V., 17, 18, 23; 68
See Rao, D.P., 11, 23, 27; 69
See Somayajulu, Y.K., 70
Satayanarayana Rao, T.S., 22; 69
Satchell, E.R., 358; 392
Saucerman, S., 410, 415, 418, 420, 490; 528
Saunders, W.B., 481, 482, 485; 528
See Hastie, L.C., 489, 496; 520
Savidge, G., 96; 103
Sayce, C.S., 172; 230
Scarabino, V. See Defeo, O., 224
Scardi, M., See Casola, E., 154
Schaan, O., 418, 434; 528
Schaefer, M.B., 429; 528
Scheibling, R.E. See Metaxas, A., 297
Scheltema, R.S., 246, 248, 249; 261
Scherman, P.A. See McQuaid, C.D., 239; 259
Schiller, C., 497; 528
Schink, T.D., 175, 176, 177, 178, 220; 230
Schneider, F.I., 141, 148; 160
Schnute, J., 493; 528
Schoeman, D., 201, 202, 203, 204; 230
Schofield, N.J. See Lukatelich, R.J., 158
Page 551
Schramm, M. See McLachlan, A., 227

Schroeder, T.E. See Lewis, C.A., 390
Schubauer, J.P., 283; 299
Schutz, C., 8; 69
Scipione, M.B. See Gambi, M.C., 155
Scott, D.A., 400; 528
Scott, H.C. See Howard, G.K., 344; 389
Scott, W.B. See Fischer, W., 225
Scotto, L.E. See Gore, R.H., 155
Scully, E.P., 278; 299
Seapy, R.R., 308; 392
Searcy Bernal, R., 169, 170; 231
Seed, R. See Williams, G.A., 280, 284, 285, 287; 302
Seeley, R.H., 240, 246, 253, 276, 287; 261, 299
Seijo, J.C., See Defeo, O., 224
Seman, R. See Adams, T.J.H., 512
Seoane, J., 1958. See Ardré, F., 384
Serchuk, F.M. See Murawski, S.A., 219; 228
Sergé, G., 98; 103
Sergeev, V.N., 137; 160
Sergievsky, S.O., 238; 261
See Zaslavskaya, N.I., 262
Sesawa, A. See Lewis, A.D., 522
See Milton, D.A., 3, 10, 22, 35, 38, 53; 69
See Rawlinson, N.J.F., 523
Sewell, R.B.S., 3, 10, 22, 35, 38, 53; 69
Shaklock, P.F., 144, 149; 160
Shaklee, J.B., 450; 528
Shapiro, L. See Maddock, J.R., 79; 102
Sharma, G.S., 50; 69
See Varadachari, V.V.R., 57; 70
Sharma, S. See Dalzell, P., 517
Sharma, S.P., 459, 460; 528
See Rawlinson, N.J.F., 458; 527
Sharp, L. See Fish, J.D., 251; 256
Sharpies, P., 410, 492; 528
Shaw, W.N., 165, 167, 168, 169, 170, 171; 231
Shay, T.S., 95; 103
Shcherbinin, A.D., 57; 69
Shelford, V.E., 306, 309; 393
Shelley, C., 474, 475; 528
Shenoi, S.S.C. See Shetye, S.R., 69
Shepherd, S.A., 118, 121; 160
See Larkum, A.W.D., 157
See Sergeev, V.N., 160
Sherwood, J.D. See Jeffery, D.J., 85; 102
Sherwood, T.S. See Hensley, R.A., 413, 415, 417; 520
Shetye, S.R., 13, 16, 21, 59, 60; 69
Shimeta, J., 82, 95; 103
Shiota, P.M., 453; 528
See Uchida, R.N., 530
Shirley, T.C., 283; 299
Shokes, R., 334; 393
Shomura, R.S., 432, 450; 528
See Polovina, J.J., 442; 525
Short, A.D., 164, 186; 231
Short, F.T., 111; 160
See Howard, R.K., 127, 141, 146, 149; 156
Shumway, S.E. See Foltz, D.W., 256
Siebens, T.L. See Drum, A.S., 225
Sieburth, J.M., 133; 160
Sienko, J. See Dippolito, A., 294
Sigurdsson, J.B., 196; 231
Silberberg, A. See Sergé, G., 98; 103
Silberstein, K., 114, 117, 118, 119, 135, 150; 160
Simons, D. See Rickard, N.A., 230
Simons, D.D. See Ayres, D.L., 173, 174, 175, 176, 177, 178, 220; 222
Simons, D.R. See Lassuy, D., 172, 173, 174, 175, 176, 177, 178, 220; 227
Simpson, C.J. See Masini, R.J., 158
Simpson, R.D., 249; 261
Sims, N., 399, 431, 447, 480, 483, 484, 485, 486; 528
Sims, N.A., 485, 486; 528
Sims, R., 203; 231
Singh, S.M., 244; 261
Siu, P. See Grand, S., 519
Skadsheim, A., 147; 160
Skewes, T., 498
Slatkin, M., 246; 261
Sloan, N.A. See Conand, C., 476; 516
Slocum, C.J., 285; 299
Smayda, T.J., 76, 83; 103
Smelcer, C. See Orth, F.L., 229
Smetacek, V. See Riebesell, U., 103
Smetacek, V.S., 83; 103
Smith, A., 406, 407, 410, 412, 419, 421, 426, 427, 504; 528
Smith, A.J., 410, 418, 420, 422, 472, 498; 529
Smith, B.D., 289, 473, 481; 299, 529
Smith, B.R. See Boy, R.L., 445; 514
Smith, B.S. See Dippolito, A., 294
Smith, D.A.S., 204, 205, 238, 251, 289, 290; 231, 261, 299
Smith, D.F. See Tranter, D.J., 161
Smith, I.D. See Hughes, R.G., 156
Smith, J. E., 240, 245; 261
Smith, M.K., 414, 440; 529
Smith, P.J., 207, 213; 231
See Richardson, J.R., 230
Smith, S.M., 235, 245; 261
Smith, T.D., 410, 506; 529
Smith Jr, W.O. See Penhale, P.A. 119, 149; 159
Smith, W.O. See Culver, M.E., 83, 101
Snyder, T.P., 245, 246; 261
Soiseth, C.R. See Dugan, J.E., 225
Somayajulu, Y.K., 24; 70
Somerton, D.A., 460; 529
Sommer, U., 76, 80, 82; 104
Song, W.C., 378, 393
Sorensen, M. See Bastida, R., 223
Soriano, M. See Gayanilo, F.C., 226
Page 552
Sornin, J.M., 147; 160

Sourie, R., 308, 310; 393
Sournia, A., 79, 81, 84, 100; 104
South, G.R., 431; 529
Southgate, T., 289; 299
Southward, A.J. See Anderson, D.T. , 339, 342, 348; 384
Southwood, T.R.E., 250, 264; 261, 299
Sparre, P., 169; 231
Spiegel, M.R., 86; 104
Spight, T.M., 250, 278; 261, 299
Springer, V.G., 404; 529
Sproston, N.G. See Moore, H.B., 276; 297
Squire, L. See Macpherson, G.R., 502; 523
Sreerama Murty, K.V. See Sundara Raman, K.V., 22; 70
SSAP, 421; 529
Stace, G., 211; 231
Stanhope, H.S., 281, 282; 299
Stanley, S.M., 188; 231
Staples, D.J., 495; 529
Stearns, S.C., 250, 251; 261
Steffensen, D.A. See Shepherd, S.A. 160
Stein, A., 410, 412, 417; 529
Steneck, R.S., 116, 119, 121, 141, 146, 284; 160, 299
See Hacker, S.D., 149; 155
StentonDozey, J.M.E., 200, 201; 231
See Brown, A.C., 223
See Matthews, S., 227
Stephenson, M.D., 167, 170; 231
See Nybakken, J., 165, 166, 168; 229
Stevenson, D.K., 428; 529
Stewart, P.C. See Kailola, P.J., 521
Stickle, W.B. See Liu, L.L., 258
See Shirley, T.C., 299
Stirling, D., 280; 299
Stiven, A.E., Stiven, A.E. See Kneib, R.T., 281; 296
Stolzenbach, K.D. See Bowen, J.D., 101
Stoner, A.W., 122, 137, 139, 140, 141, 142, 148, 149; 160
See Lewis, F.G., 148; 157
Stotter, D.R. See Cranfield, H.J., 223, 224
Strathmann, R.R., 249; 261
See Palmer, A.R., 249; 260
Straughan, D., 361, 366, 380; 393
Street, R.J., 207, 209, 211; 231
Strickland, D.H. See Eppley, R.W., 101
Strong, K.W., 143; 260
Struhsaker, J.W., 242; 261
Struhsaker, P., 460; 529
Stubbings, H.G., 308, 310; 393
Sua, D., 440; 529
Sua, T. See Zann, L.P., 531
Sudekum, A.E., 425, 451; 529
Sullivan, M.J., 132, 135; 160
Summerson, H.C. See Peterson, C.H., 249; 260
Sumpton, W. See Brown, I.W., 575
Sundar, D. See Shetye, S.R., 69
Sundara Raman, K.V., 22; 70
Sundberg, P., 240, 438; 261, 529
See Janson, K., 240; 257
See Johanneson, K., 235; 258
See Richards, A.H., 418, 438, 439; 527
Sundell, K., 240; 261
Sundstrom, B., 13; 70
Suryanarayana, A., 11, 15, 19, 21, 22, 23, 27, 37, 52, 59; 70
See Varkey, M.J., 1–70
Sutherland, J.P. See Menge, B.A., 272; 297
Sverdrup, H.U., 53, 54; 70
Swadling, P., 478, 479, 480; 529
Swallow, J.C. See Cutler, A.N., 56; 67
Swamy, K., 487, 488, 492, 494; 529
Swenson, R.P. See Gilly, W.F., 280; 295
Sze, P., 273, 275; 299
Szelistowski, W.A. See Borjesson, D.L., 282; 293
Tabe, T. See Tebano, T.N., 453, 457; 529

Tagami, D.T. See Ralston, S., 469; 526
See Uchida, R.N., 530
Talavero, F., 206; 231
Talbot, F.H. See Goldman, B., 426; 519
Talbot, M.M.B., 248; 261
Talikhedkar, P.M., 205; 231
See Nagabhushanam, R., 205, 206; 228
Taman, J. See Adams, T.J.H., 512
Taniera, T. See Mees, C.C., 523
Tankersley, R.A., 280; 299
Tarazona, J. See Arntz, W.E., 222
Tarifeño, E., 191, 192; 231
Tarr, R., 202, 203; 231
Tatarenkov, A., 239; 261
Tatarenkov, A.N., 239, 247; 262
See Zaslavskaya, N.I., 262
Tateno, Y. See Nei, M., 243; 259
Taumaia, P., 410, 418, 449, 457; 529
Taylor, G.I., 93; 104
Taylor, R. See Coyne, T., 516
Taylor, T.D. See Acrivos, A., 78; 101
Teal, J.M., 282; 300
See Connor, M.S., 294
Tebano, T.N., 453, 457, 482; 529
Tegelberg, H.C., 173, 174, 178; 231
Tegner, M.J., 142; 160
Teissier, G. See Prenant, M., 307; 392
Temkin, M.H., 280, 281, 283; 300
See Stanhope, H.S., 299
Tenakanai, C.D., 495; 530
Tenekanai, C.T. See Opnai, L.J., 465; 525
Tennekes, H., 92, 93; 104
Tenore, K.R., 147; 161
Page 553
Teopa, M. See Nash, W., 524

Terekia, O. See Nash, W., 524
Teulieres, M.H. See Leblic, I., 410; 522
Tevkolsky, S.A. See Press, W.H., 103
Thayer, G.W. See Zieman, J.C., 162
Theede, H. See AbeleOeschger, D., 288; 293
Theisen, D. See Coates, D., 516
Thirupad, P.U. See Varadachari, V.V.R., 70
Thollot, P., 404, 466, 467, 470, 471; 530
Thoman, T.A. See Heck, K.L., 149; 155
Thomas, C.D. See Sieburth, J.M., 133; 160
Thomas, H.J., 317, 318; 320, 323, 341, 393
Thomas, M.L.H., 289, 290; 300
See Rangeley, R.W., 272, 287; 298
Thomas, W.H., 97; 104
Thompson, P.A. See Dortch, Q., 101
See Waite, A.M., 104
Thorpe, J.P. See Moyse, J., 259
Thorson, G., 249; 262
Throndsen, J., 79, 81; 104
Tilves, F. See Molares, J., 391
Tirard, P. See Kulbicki, M., 521
Tiroba, G., 462; 530
See Blaber, S.J. M., 514
Titman, C.W. See Sigurdsson, J.B., 231
Titman, D., 83; 104
Tobias, W.J. See Tsuda, R.T., 530
Tomlinson, P.K., 168, 169, 171; 231
Torres Jr, F. See AriasGonzalez, J.E., 513
Towler, E.D., 306; 393
See Shelford, V.E., 306; 393
Tranter, D.J., 138; 161
Tranter, H.A. See Tranter, D.J., 161
Trautman, D.A., 134; 161
Trocine, R.P., 119; 161
Troll, C. See Landsberg, H.E., 67
Trowbridge, C.D., 274, 289; 300
Trueman, E.R., 183, 201; 231
See Ansell, A.D., 183; 222
See Brown, A.C., 223
Trussell, G.C., 240; 262
Tsuda, R.T., 424; 530
Tuara, P. See Nash, W., 524
Tu’avao, T., 412; 530
Tufts, D.F. See Sayce, C.S., 172; 230
Tulua, S. See Latu, S., 433, 439, 441, 503, 506; 522
Tumaia, P. See Preston, G.L., 526
Tuqiri, S. See South, G.R., 529
Turgeon, K.W. See Fralick, R.A. F., 295
Turk, S.M., 307; 393
Turner, L. See Berg, H.C., 79; 101
Uchida, R.N., 451, 491, 493; 530

Uchiyama, J.H. See Struhsaker, P., 460; 529
See Uchida, R.N., 451, 493; 530
Udagawa, K. See Tu’avao, T., 530
Ugarte, R. See Santelices, B., 141, 146; 160
Ulaiwai, W.K., 465, 467, 471; 530
Underwood, A.J., 237, 247, 250, 265, 267, 268, 269, 270, 274, 276, 277, 279, 284, 289, 291, 292; 262, 300
See Chapman, M.G., 268; 294
See Creese, R.G., 237; 255
See Fairweather, P.G., 267, 268, 269; 295
See Moran, M.J.; 297
UNESCO, 7; 70
Unnikrishnan, A.S. 61; 70
See Bahulayan, N., 61; 66
Unny, M.M. See Chari, S.T., 206; 223
Ursin, E. See Sparre, P., 231
Uwate, R., 410, 418; 530
Uysal, H. See Geldiay, R., 195; 226
Vadas, R.L., 145, 271, 272, 273, 274, 279, 291; 161, 300, 301
See Miller, S.L., 274; 297
Vakamoce, V. See Preston, G.L., 526
Vakily, J.M., 169; 231
Valdevenito, I. See Peredo, S., 229
Valdivieso, V. See Defeo, O., 224
Valentine, J.F., 119, 145; 161
Valiela, I. See Connor, M.S., 294
Van Alstyne, K.L., 286; 301
See Boulding, E.G., 240, 252; 255
Van Blaricom, G.R. See Estes, J.A., 168, 173, 175, 176, 177; 225
Vance, R.R., 282; 301
Van den Ben, D., 117, 161
van der Horst, G., 200; 231
See McLachlan, A., 227
Van de Ven, T.G.M., 95; 104
Van Duuren, F., 91; 104
van Geel, G. See Jacobs, R.P.W.M., 156
Van Ireland, E.T., 81; 104
See Nienhuis, P.H., 122; 159
Van Marion, P., 240; 262
Van Montfrans, J., 109, 116, 118, 119, 123, 124, 125, 136, 141, 142, 144, 146, 149; 161
See Orth, R.J., 109, 110, 117, 119, 141, 147; 159
Van Sickle, J., 182; 232
Van Valen, L., 244; 262
Varadachari, V.V. R., 10, 23, 52, 57; 70
See Bahulayan, N., 56; 66
See Murty, C.S., 19; 68
Varkey, M.J., 1–70; 8, 23, 35, 38, 50, 51, 52, 53, 54, 55, 56, 58, 59, 60, 61, 62, 70
Varma, P.V. See Murty, C.S., 10; 68
Vaughn, C.C., 280, 281; 301
Vay, S.A. See Van Montfrans, J., 161
Veit, R.R. See Helmuth, B., 256
Velez, A., 183, 184, 185, 186; 232
Venables, B.J. See Vélez, A., 232
Venema, S.C. See Sparre, P., 231
Venkataswaran, S.V., 8; 70
Page 554
Vermaat, J.E. See Hootsmans, M.J., 118, 122, 126, 141, 143, 149; 156
Vermeij, G.J., 237, 240, 242, 246, 249, 264, 271, 276, 279, 262; 301
See Blundon, J.A., 282; 293
Veron, J.E.N., 400; 530
Veronis, G., 56; 70
Verploegh, G., 24; 70
Vetterling, W.T. See Press, W.H., 103
Viala, F. See Preston, G.L., 526
Viejo, R.M., 287; 301
Vinci, D. See Russo, G.F., 159
Virnstein, R.W., 116, 117, 136, 137, 138, 148; 161
See Kulczycki, G.R., 157
See Nelson, W.G., 158
Viswambharan, N.K., 24; 70
Vivas, A.M. See Gaines, M.S., 256
Von Bertalanffy, L., 189; 232
Vuilleumier, F., 236; 262
Wade, B.A., 182, 183, 184, 185; 232

Waffy, A., 495; 530
Wagner, F.J. E., 271; 301
Waite, A.M., 83; 104
Waite, M.E., 329, 332, 333; 393
Walker, D.I., 111, 114, 115, 117, 133; 161
See Brearley, A., 122; 153
See Harlin, M.M., 155
See Kendrick, G.A., 156
See Kirkman, H., 111; 157
Walker, G., 304, 321, 326, 329, 332, 334, 348, 369, 370, 373; 393
See Waite, M.E., 329, 332, 333; 393
See White, K.N., 334; 393
See Yule, A.B., 321; 394
Walker, M.H., 423, 502; 530
Walley, L.J., 373, 374; 375, 393
Walter, H., 8; 70
Walters, C.K., 423; 530
Walters, C.J. See Hilborn, R., 188; 226
See Hilborn, R., 501, 502; 520
Wankowski, J.W.J., 432; 530
Wantiez, L., 466, 471; 530
See Kulbicki, M., 466, 467, 470; 521
Ward, R.D., 236, 239, 244, 246, 247, 248; 262
See Janson, K., 236, 239, 244; 257
See Knight, A.J., 236, 246; 258
Ward, R.G., 473; 530
Ward, S., 268; 301
Ward, T.J., 138; 161
Ware, G.N. See Lewin, J., 227
Warmoes, T. See Backeljau, T., 236; 254
See Johanneson, K., 244, 245, 249; 258
Warner, R.W., 167; 232
Warpeha, P.R. See Quinn, N., 526
Warren, J.H., 281, 282; 301
Warwick, T., 236; 262
See Atkinson, W.D., 237, 238; 254
See Ward, R.D., 236, 246; 262
Wass, R.C., 407, 427, 428, 430; 530
Watling, L. See Steneck, R.S., 116, 119, 121, 141, 284; 160, 299
Watson, D.C., 273, 274, 276, 284, 285, 286, 287; 301
See Hawkins, S.J., 295
See Norton T.A., 298
Watson, R.A., 455, 464, 470, 471, 472; 530
Watt, P., 413, 452; 530
Wauthy, B., 402; 530
Wear, R.G., 209, 213; 232
Weatherall, J.A., 460; 530
Weber, M.E. See Clift, R., 101
Weese, A.O. See Shelford, V.E., 393
Wells, F.E., 136; 161
Wells, G.N. See Trocine, R.P., 161
Wells, S.M., 400, 401; 531
Weltner, W., 305, 306, 308; 393
Wendell, F., 167, 168, 170, 171, 174; 232
West, D.L., 281; 301
West. L., 267, 379; 301, 393
West, R. See Shepherd, S.A. 160
West, R.J., 111; 161
See Larkum, A.W.D., 120; 157
Westoby, M. See Bell, J.D., 122, 135, 149; 153
Wetherbee, B.M. See Holland, K.N., 520
Wethey, D.S., 292; 301
Wetstone, G.S. See Heck, K.L., 140; 155
Wetzel, R.L., 119; 161
See Neckles, H.A., 158
See Van Montfrans, J., 161
Weymouth, F.W., 172, 173, 174; 232
White, K.N., 334; 393
White, T.J. See Wilson, A.C., 262
Whitehead, P.J.P., 453; 531
Whitford, J. See Nash, W., 524
Wilber, T.P., 278, 281, 282; 301
Wilhelmsen, U., 271, 273; 301
Wilkins, N.P., 235, 247; 262
Willcocks, P.A., 133; 161
Williams, A.B., 138; 161
Williams, A.H. See West, D.L., 281; 301
See McBride, C.J., 297
Williams, D.M., 426; 531
See Munro, J.L., 422; 524
Williams, G.A., 265, 269, 280, 284, 285, 287, 292; 301, 302
See Norton, T.A., 298
Williams, H.A. See Ralston, S., 423, 424, 440; 526
Williams, J. See Kulbicki, M., 521
See Orth, F.L., 229
Williams, M.J. See Kailola, P.J., 521
Williams, R., 339, 355, 357; 394
Williams, S.L., 110, 122, 130, 141, 149, 151; 161
Page 555
Wilson, A.C., 243; 262

Wilson, A.M., 410; 531
Wilson, E.O. See MacArthur, R.H., 249; 258
Wilson, K.A. See Gore, R.H.; 155
Wilson, M.A., 451; 531
See Kailola, P.J., 404, 455, 466, 467; 521
Winet, H. See Brennen, C., 83; 101
Winter, M.A. See Hamilton, P.V., 280; 295
Winterbottom, R. See Emery, A.R., 410, 411, 419; 518
Withers, T.H., 304; 394
WiumAndersen, G., 245, 247; 262
WiumAndersen, S. See Borum, J., 133; 153
Woelkerling, W.J. See Bramwell, M.D., 116, 117, 133, 134; 153
See Bulthuis, D.A., 111, 114, 117, 118, 133, 134, 135; 153, 154
See Harlin, M.M., 155
See Walker, D.I., 117, 133; 161
WolfGladrow, D.A. See Riebesell, U., 103
Womersley, H.B.S., 134; 162
Wongratana, T. See Whitehead, P.J.P., 531
Wood, E.F.J., 135; 162
Woodroffe, C.D., 400; 531
Woodward, B.B., 250; 262
Wooldridge, T. See McLachlan, A., 227
Wright, A., 399, 410, 418, 419, 421, 422, 424, 428, 435, 446, 449, 479, 480, 483, 484, 500; 531
See Dalzell, P., 410, 411, 428, 449; 517
See Yeeting, B., 410, 413, 429, 431, 504; 531
Wright, L.D., See Short, A.D., 164, 186; 231
Wrobel, L., 434; 531
Wu, T.Y. See Chwang, A.T., 83; 101
See Keller, S.R., 84; 102
Wyrtki, K., 3, 22, 23, 25, 43, 52, 60; 70
Yager, P.L. See Jumars, P.A., 102

Yaglom, A.M. See Monin, A.S., 92; 103
Yamaguchi, M., 479; 531
Yamaguchi, Y., 460; 531
Yamazaki, H., 95; 104
Yang, J. See Yu, L., 70
Yasugi, R., 372; 394
Yates, J.L., 286; 302
Yeeting, B., 410, 413, 429, 431, 504; 531
Yeeting, B.M. See Mees, C.C., 453; 523
See Sims, N.A., 528
Yen, S. See Grand, S., 519
Yesaki, M., 11, 21; 70
Yoshimoto, S.S. See Clarke, R., 491, 493; 516
Yoshino, T., 361; 394
Young, D.K., 122, 142; 162
Young, M.W. See Young, D.K., 122, 142; 162
Young, N. See McLachlan, A., 201; 227
Young, P.C., 136, 140; 162
See Kirkman, H. 120, 145; 157
See Ward, T.J., 138; 161
Younglao, D. See Hunte, W., 249; 257
Youngson, A. See Walker, G., 321; 393
Yu, L., 61; 70
Yule, A.B., 321; 394
See Cardosa, A.C., 361, 362, 366; 386
See Kugele, M., 356, 357; 390
Yund, P.O. See Bertness, M.D., 293
Yuriar, F.J.L. See Searcy Bernal, R., 231
Zaklinskii, G.B., 43; 70

Zann, L.P., 410, 412, 428, 484, 487, 493, 498; 531
Zapata, O., 118; 162
Zaslavskaya, N.I., 246; 262
Zevina, G.B., 304; 394
Zieman, J.C., 114; 162
Zieman, R.T. See Zieman, J.C., 162
Zimmerman, R.J., 119, 120, 122, 123, 124, 136, 141, 142, 143; 162
Zingmark, R. See Foster, M., 388
Zouros, E., 244; 262
See Mallet, A.L., 258
Zoutendyk, D., 472, 473, 498; 531
Zupo, V. See Gambi, M.C., 155
Page 556

Page 557
SYSTEMATIC INDEX
UCL Press
References to complete articles are given in bold type; references to sections of articles are given in italics; references to pages are given in normal type.
Acanthocybium solandri, 406, 407, 447, 449

Acanthophora, 116
Acanthopleura gemmata, 481
Acanthuridae, 401, 404, 415, 416, 417, 418, 419, 420, 421, 426, 431
Acanthurus bleekeri, 407
dussumieri, 415
lineatus, 423
nigricauda, 423
nigrofuscus, 423
triostegus, 423
xanthopterus, 407, 423
Acmaea digitalis, 309
Actinopyga, 404, 474
echinites, 473, 474, 475, 476, 477
leconora, 474
Mauritania, 473, 474, 476, 477
miliaris, 473, 474, 475, 477
Albula glossodonta, 504
Albulidae, 416, 420
Allorchestes compressa, 143
Amblygaster, 456, 461
clupeoides, 452, 460
sirm, 452, 456, 457, 460, 462
Amphibolis, 111, 115, 118, 119, 120, 130, 133, 136, 137, 138, 139, 140, 142
antarctica, 112, 114, 115, 118, 133, 134, 135, 137, 147, 148
griffithii, 111, 112, 115, 121, 128, 130, 135, 137, 139, 142, 147, 148
Amphiroa, 116
fragilissima, 120
Amphora, 125, 146
Ampithoe, 129, 130, 138, 143, 148
longimani, 143
marcuzii, 143
mea, 143
Amusium pleuronectes, 480
Anachis, 126
avara, 139
avara sempilicata, 126
Anadara, 481
antiquata, 480, 481
cornea, 481, 482
granosa, 480, 481
maculosa, 479, 482
Anaulus australis, 201, 248
Anguilla obscura, 470
Anodontia edentula, 481
philippina, 480
Aphareus furca, 436
rutilans, 435, 436, 438
Aphriza virgata, 379
Aplysia, 121
Apogonidae, 421, 469
Aprion virescens, 436, 440
Apsilinae, 435, 436
Araneus cribrarius, 184
Area ventricosa, 481, 482, 483
Archamia, 456
zosterophora, 460
Arctica islandica, 219
Ariidae, 404, 467, 468
Artemia, 345
salina, 345
Asaphis violascens, 480, 481
Ascidiacea
Ascophyllum nodosum, 143, 273, 274, 279, 284, 286, 287, 291
Assiminea californica, 279
Asterionella, 85
glacialis, 187
Astyris lunata, 127
Atactodea striata, 480, 481
Atherinidae, 408, 421, 452, 455, 469
Atherinomorus, 456
lacunosus, 456, 460
Audouinella, 133
Austrocochlea constricta, 270
Auxis, 447
rochei, 451
thazard, 449, 451
Avicennia, 400
Balanus balanus, 367

eburneus, 322
glandula, 308, 309, 334, 338, 339, 357, 358
nubilus, 322
perforatus, 310, 367, 370
tintinnabulum, 310
Balistidae, 416, 417, 419, 431
Barbitia reeveana, 481
Page 558
Batissa fortis, 481

violacea, 479, 41, 482
Belonidae, 407, 416, 417, 446, 448
Bembicium, 235, 269
nanum, 269–71; 269, 270, 271, 277
vittatum, 248
Birgus latro, 404, 489
Bittium, 126, 136, 138, 146
varium, 125, 126, 127, 129, 139, 144, 146
Blenniidae, 431
Blennius pholis, 238
Bohadschia, 404, 472
argus, 474
marmorata, 474
vitiensis, 474
Bolbometopon muricatum, 413, 415, 417, 424, 425, 431, 504
Bothidae, 469
Bruguiera, 400
Bryothamnion, 116
Bryozoa, 116
Buccinanops duartei, 188
Bulla striata, 127
Caecum, 136
Caesio, 456
Caesionidae, 408, 420, 427, 452
Calantica, 304, 329
spinosa, 304, 323, 329, 334, 349, 361, 372
Callinectes, 126
exasperatus, 184
sapidus, 281
Cancer magister, 173
Cancer pagurus, 272
Cantharidus bellulus, 137
irisodontes, 137
lepidus, 131, 137, 140, 145
pulcherrimus, 137
Capitulum, 304
mitella, 329, 361, 369, 372
Caprella laeviuscula, 124, 150
Carangidae, 404, 408, 413, 416, 417, 418, 420, 421, 425, 432, 435, 439, 448, 452, 453, 468, 470
Caranx, hippos, 181
ignobilis, 425, 450, 451, 466
lugubris, 435, 441
melampygus, 425, 450, 451
papuensis, 450
Carcharhinidae, 417
Carcinus, 287
maenas, 240, 242, 272, 275, 287
Cardisoma, 488
carniflex, 404
Cardium fragum, 481, 482, 483
Carpilius maculatus, 404, 488
Cathorops spixii, 181
Cellana tramoserica, 270
Centropomidae, 404, 468
Cepaea, 237
Cephalopholis aurantia, 436
Ceramiaceae, 133
Ceramiales, 133
Cerapus abditus, 137
Cercaria caribbea XLII, 184
Cerithium muscarum, 127, 139
tenellum, 128, 145
Chaceon, 489
granulatus, 489
Chaetoceros armatum, 173
Chaetodon miliaris, 425
Chaetodontidae, 431
Chaetomorpha, 274
Chama, 481
isotoma, 481
sordida, 481
Chamelea gallina, 199
Chanidae, 416, 417, 420, 421, 468
Chanos chanos, 466
Cheilinus undulatus, 413, 417, 425, 431
Cheilopogon, 455
antoncichi, 455
atrisignis, 455
spilonopterus, 455
suttoni, 455
unicolor, 455
Chlorophyta, 134
Chondria, 116, 133
Chondrus crispus, 144, 273, 274, 285, 287, 289
Chordata, 116
Chromis albipectoralis, 431
Chrysiptera cyanea, 431
Chrysophrys auratus, 209
Chthamalus, 334, 349, 353, 357
dalli, 308, 309, 339
dentatus, 310
fissus, 309, 356
montagui, 352
stellatus, 310, 352
Cirrhitidae, 404, 431
Cirripedia, 303, 304
Cladophora, 116, 127, 143, 145
ruchingeri, 143
Clupeidae, 404, 408, 420, 421, 452, 468
Clytia bakeri, 167
Cnidaria, 116
Coccolithus huxleyi, 76
Cocconeis scutellum, 125, 142, 146
Codakia punctata, 480, 481
tigerina, 481
Codium, 143, 310
Coleochaete, 145
Cololabis saira, 433
Conchocelis, 378
Conchoderma auritum, 306
Concholepas concholepas, 194
Conodon nobilis, 184
Page 559
Corallina, 116, 310

Coronula diadema, 306
Corophium acutum, 136
ascherusicum, 136
insidiosum, 136, 148
Coryphaena hippurus, 407, 446, 449, 450, 451
Coryphaenidae, 448
Coscinodiscus, 187
wailesii, 76
Crassostrea rhizophorae, 481
Crenimugil crenilabis, 417
Crepidula, 136
convexa, 139
Crocethia alba, 184, 205
Crustacea, 303
Ctena bella, 481
Ctenochaetus binotatus, 423
striatus, 423
tomiensis, 431
Cymadusa, 126, 138
compta, 124, 127, 136, 139, 143
Cymodocea, 111
Cymodoceaceae, 112
Cymodocea filiforme, 112
serrulata, 112, 145
Cyprinus carpio, 466, 500
Cyproidea, 138, 148
Cypselurus, 455
angusticeps, 455
oligolepis, 455
pitcairnensis, 455
poecilopterus, 455
Dascyllus aruanus, 431

Decapterus, 456, 461
macarellus, 453, 455, 460
macrosoma, 456, 460
russelli, 460
Delesseria sanguinea, 290
Dermatolithon, 133
Diadema setosum, 473
Diagramma pictum, 424
Dictyosiphon, 144
Dictyota dichotoma
Dinophysis acuta, 76
Ditylum brightwelii, 91
Donacidae, 165
Donax, 182, 183, 184, 186, 195, 196, 199, 200, 205
cuneatus, 163, 164, 165, 168, 204, 205, 206, 217, 221
deltoides, 163, 164, 165, 168, 206, 207, 217, 218
denticulatus, 163, 164, 165, 168, 180, 182, 183, 184, 185, 186, 217, 221
faba, 163, 164, 165, 168, 204, 205, 206, 217, 221
gouldi, 168
hanleyanus, 188, 218
semistriatus, 195
serra, 163, 164, 165, 168, 199, 200, 201, 202, 203, 204, 206, 217, 218, 220, 221, 253
sordidus, 199
striatus, 163, 164, 165, 168, 180, 182, 183, 184, 185, 186, 217, 221
trunculus, 163, 164, 165, 168, 194, 195, 196, 197, 198, 199, 217, 221
venustus, 195
vittatus, 195, 196, 197
Dussumieria, 452, 460, 461
Eatoniella australiensis, 139

Echinoidea, 472
Echinometra mathaei, 473
Ecklonia radiata, 143
Ectocarpus, 116, 143
Elagatis bipinnulatus, 445, 449, 455
Eleotridae, 468
Emplectonema gracile, 379
Encrasicholina, 408, 454, 456, 462
devisi, 452, 456, 458, 460, 461, 462, 503
heteroloba, 452, 456, 458, 460, 461, 462, 503
punctifer, 456, 460
purpurea, 454, 459
Engraulidae, 404, 408, 452, 468
Enhalus, 111, 118
acoroides, 112, 114, 128, 133, 145
Enhydra lutris, 167, 173
Enosania olevata, 481
Ensis minor, 199
Enteromorpha, 143, 144, 146, 269, 271, 272
intestinalis, 273, 275
linziformis, 143
Epinephelus areolatus, 425, 436, 437
chlorostigma, 436, 437
cometae, 436, 437
fasciatus, 414, 425, 436
flavocaeruleus, 436
hoedti, 425
maculatus, 425
magniscutis, 436
merra, 425
miliaris, 436, 437
morrhua, 436, 437, 439, 441, 442
polyphekiadon (=microdon), 425
retouti, 436, 437
septemfasciatus, 436, 437, 439, 441, 442
tauvina, 425
Erichsoniella attenuata, 127, 129
Ericthonius, 137
Escherichia coli, 79
Etelinae, 435, 436
Etelis, 432, 437
carbunculus, 436, 437, 438, 439, 440, 441, 442
coruscans, 433, 435, 436, 438, 439, 440, 441, 442, 443
radiosus, 435, 436
Etisus splendidus, 404, 488
Eucinostomus, 181
Eugymnanthea cirrhifera, 181
Page 560
Eunice viridis, 405

Euthynnus affinis, 445, 447, 449, 451
Excirolana armata, 188
Exocoetidae, 461
Exocoetus, 455
Fimbriata fimbriata, 481

Fucus, 114, 271, 273, 274, 284, 286, 287, 288, 289, 307, 310
ceranoides, 273
distichus, 275, 286, 288
edentatus, 290
evanescent, 275, 288
serratus, 238, 274, 284, 287, 290
spiralis, 288
vesiculosus, 273, 274, 275, 284, 286, 287, 288
Fundulus heteroclitus, 281
Gafrarium pectinatum, 481

tumidum, 480, 481
Gaimardia, 249
Gammarus, 129, 130
mucronatus, 124, 127, 143, 148
Gastropoda, 181
Gazza minuta, 460
Geloina coaxans, 480
Gempylidae, 435
Gerridae, 416, 417, 467, 469
Gibbula, 272
ardens, 128
umbilicaris, 127, 128, 144, 146, 150
Giffordia mitchelliae, 148, 149
Gigartina papillata, 286
Gitanopsis, 138, 148
Gleotrichia, 145
Glycera, 181
Gobiidae, 431
Gonyaulax grindleyi, 203
Gracilaria, 136, 148
Grammatorcynus bicarinatus, 447, 450
Grandidierella bonnieroides, 124, 127, 143
Gymnocranius, 423
japonicus, 423, 436
lethrinoides, 423
rivulatus, 423
Gymnosarda unicolor, 415, 450
Gyrodinium dorsum, 76
Haematopus ostralegus, 188

Haemulidae, 416, 417, 419, 424, 467, 469
Halodule, 111, 137, 139, 142, 148
beaudetti, 112
uninervis, 112
wrightii, 112, 114, 124, 126, 127, 136, 138, 139, 140, 143, 144, 146, 148, 150
Halophila, 111
ovalis, 113, 114, 115, 118
Harpontidae, 467, 468
Hemicentrotus pulcherrimus, 473
Hemiramphidae, 416, 417, 446, 453, 461, 469
Herklotsichthys, 452, 456
quadrimaculatus, 453, 456, 460, 461
Heterocentrotus mammilatus, 473
Heteroderma, 133
lejolisii, 126
Heterozostera, 111
tasmanica, 111, 114, 115, 118, 125, 133, 134, 142
Hildenbrandia prototypus, 270
Hippolyte, 139
pleuracanthus, 127
Hippopus hippopus, 480, 481, 483, 484
Hipposcarus longiceps, 415
Hirundichthys, 455
Holocentridae, 404, 416, 418, 419, 420
Holothuria, 475
atra, 473, 474, 476, 477
cinarescens, 473
edulis, 474
fuscogilva, 474, 475, 477
fuscopunctata, 474
leucospilata, 473
nobilis, 474, 475, 477
scabra, 472, 473, 474, 475, 476, 477
verrucosa, 473
Holothuroidea, 472
Hormosira banksii, 270
Hyale, 138, 148
rubra, 143
Hydrobia, 126, 127, 144, 276
ulvae, 126, 144, 276, 277
Hydrocharitaceae, 112
Hydrozoa, 116
Hypnea, 143, 144, 145, 149
Hypoatherina, 456
ovalau, 455, 456, 460
Ibla, 334
cumingi, 322, 323
Idotea, 126, 127, 130, 150
baltica, 127, 129, 130, 143, 144
chelipes, 122, 126, 144
granulosa, 144
resecata, 130, 149
viridis, 127, 144
Ilyanassa (Nassarius) obsoleta, 277
obsoleta, 277, 278
Iridaea cordata, data, 290
Isocheles wurdemanni, 181
Isochrysis, 372
galbana, 372
Istiophoridae, 447, 448
Istiophorus platypterus, 449
Jububinus striatus, 127, 144
Katsuwonus pelamis, 397, 447, 449

Page 561
Kuhlia rupestris, 466, 470, 471, 472

Kyphosidae, 417, 419, 421
Kyphosus bigibbus, 414
Labridae, 413, 416, 417, 419, 420, 421, 425, 431, 439
Lactaridae, 469
Lacuna, 289–90; 138, 234, 235, 249, 251, 286, 289, 290, 291
marmorata, 290
pallidula, 247, 251, 284, 289, 290
parva, 247, 289, 290
variegata, 290
vincta, 247, 251, 288, 289, 290
Lacuninae, 234, 235, 242
Laevilitorininae, 234, 235, 242
Lambis chiragra, 481
lambis, 480, 481
truncata, 481
Laminaria, 290
digitata, 290
longicruris, 290
saccharina, 290
Larus glaucescens, 379
occidentalis, 379
Lates calcarifer, 464
Latreutes, 139
Laurencia poitei, 148
Leiognathidae, 404, 468
Leognathus bindus, 460
Lepas, 321, 325, 326, 334, 353
anatifera, 329, 370
fascicularis, 321, 353
Lepsiella vinosa, 367, 368
Lethrinidae, 404, 411, 413, 415, 416, 147, 418, 419, 420, 421, 426, 432, 435, 439, 469
Lethrinus, 423
chrysostomus, 414, 436, 439
harak, 423
kallopterus, 436, 437
mahsena, 423
miniatus, 415, 423, 436, 437, 438
miniatus (=chrysostomus), 423, 437
nebulosus, 411,415, 423
nematacanthus, 423
olivaceus, 423, 437
olivaceus (=miniatus), 423, 437
ramak, 415
reticulatus, 436
rubrioperculatus, 423
semicinctus, 423
variegatus, 436, 437
Limnoriidae, 122
Lintricula auricularia, 181
Lithotrya, 322, 327, 328, 339, 368
Littoraria, 234, 235, 238, 244, 249, 264
aberrans, 249
albicans, 249
angulifera, 247, 249
filosa, 238, 239
irrorata, 280–83; 263, 264, 276, 278, 280, 281, 282, 283, 291
pallescens, 238, 239
scabra, 252
Littorina, 241; 126, 127, 235, 238, 240, 241, 242, 244, 246, 251, 265, 271, 283, 289
angulifera, 247
arcana, 235, 236, 237, 239, 245, 246, 252
brevicula, 247
fasciata, 282
kurila, 241
littoralis, 235, 284, 287
littorea, 241–2, 271–9; 126, 142, 144, 234, 235, 236, 239, 240, 241, 242, 244, 246, 247, 248, 249, 250, 264, 265, 266, 269, 270, 271, 272, 273, 274, 275, 276, 277, 278, 279, 286, 287, 288, 291
mandshurica, 247
mariae, 235, 236, 237, 238, 246, 250, 284, 287
neglecta, 125, 139, 140, 147, 150, 235, 239, 251, 276, 283
neritoides, 244
nigrolineata, 235, 239, 240, 246
obtusata, 287–9; 235, 236, 240, 246, 265, 272, 276, 284, 286, 287, 288, 291
peruviana, 288
picta, 242
plena, 276
punctata, 244
rudis, 237, 242, 247, 251
saxatilis, 239–40; 127, 144, 235, 236, 237, 238, 239, 240, 242, 244, 245, 246, 247, 248, 249, 250, 251, 252, 264, 271, 277, 279, 288
scabra, 235, 480
scutulata, 265, 274, 285, 286
sitkana, 241, 286
squalida, 238
subrotundata, 279
unifasciata, 267
varia, 282
Littorinidae, 233–62, 263–302; 234, 252
Littorininae, 235, 242, 252
Liza vagiensis, 455
Lucina corrugata, 480
Lumbriconereis, 181
Lutjanidae, 404, 413, 415, 416, 417, 418, 419, 420, 421, 424, 426, 432, 435, 439, 448, 468
Lutjaninae, 435
Lutjanus, 424
amabilis, 424
argentimaculatus, 435, 436, 437, 438, 466, 470
bohar, 424, 435, 436, 437, 438
erythopterus, 424, 438
fulviflamma, 424
fulvus, 424
gibbus, 415, 424, 435, 436, 437
Page 562
goldei, 472
johnii, 472
kasmira, 424, 436, 437
malabaricus, 424, 436, 437, 438, 439
monostigma, 436
quinquelineatus, 424
rufolineatus, 436
sebae, 424
vitta, 424
Lynseia, 122
Lytechinus variegatus, 145
Macrobrachium, 464, 489, 497

rosenbergii, 489
Mactracea, 165, 188
Makaira mazara, 449, 450
Malacobdella grossa, 172
Marginellidae, 181
Mastogloia, 142
Megalopidae, 469
Megalops cyprinoides, 466, 471
Melarhaphe cincta, 252
neritoides, 244, 246, 248
scabra, 252
Melita nitida, 124, 143
Melongea corona, 281
Menticirrhus littoralis, 181, 184
martinicensis, 181
Mesodesma donacium, 163, 164, 165, 168, 191, 192, 193, 217, 219, 220, 221
mactroides, 163, 164, 165, 168, 186, 187, 188, 190, 199, 217, 218, 221
Mesodesmatidae, 165
Metagoniolithon, 116
Metapenaeus, 495
affinis, 490
anchistus, 405, 491
demani, 405, 490, 493
dobsoni, 490
elegans, 491
ensis, 405, 488, 490, 491, 496
Microdeutopus damnoniensisi, 136
gryllotalpa, 136
Microdiscula charopa, 139
Micropogonias furnieri, 188
Mitella, 304
Mitra mitra, 481
Mitrella lunata, 129, 148
Modiolus auriculatus, 481
philippinarum, 481
Modulus modulus, 124, 127, 139
Monodactylidae, 469
Monodonta lineata, 272
Morula, 268, 269
marginalba, 268, 269
Mugilidae, 404, 416, 417, 420, 421, 453, 466, 468
Mullidae, 413, 416, 417, 420, 421, 468
Mulloides flavolineatus, 407, 417, 425
Muraenidae, 469
Mustelus schmiti, 188
Mylobatidae, 417
Myripristis amaena, 425
Mytilus, 308, 309, 383, 384
africanus, 310
californianus, 308, 309, 380
edulis, 238, 248, 310
Naso, 415
brevirostris, 415, 423
unicornis, 415, 423
Natica cayenensis, 181
Nautilus belauensis, 481, 485
pompilius, 481, 485
Nemipteridae, 423, 467, 469
Nemipterus peroni (=furcosus), 423
Neotia ponderosa, 481
Nerita, 480, 481
atramentosa, 270, 277
maxima, 481
ornata, 481
peloronta, 481
plicata, 481
polita, 481
undata, 481
Nitzschia, 125, 142, 146
Nodilittorina, 235, 264, 269
africana, 238, 239
angustior, 239
cincta, 252
hawaiiensis, 242
lineolata, 239
punctata, 244
pyramidalis, 268
rugosa, 268
unifasciata, 267, 268, 269, 274, 287
Notoacmea testudinalis, 277
Nucella, 240–47; 241, 242, 379
emarginata, 267, 379
lamellosa, 241, 379
lapillus, 239, 240, 241, 274
Nucula rugosa, 481
Octopus cyanea, 481

Ocypode albicans, 184
ceratophthalma, 205
quadrata, 184
Olivancillaria vesica auricularia, 188
Olivella verreauxi, 184
Olividae, 181
Onchorhynchus mykiss, 466
Onuphis, 207
Ophioscion punctatissimus, 181
Oreochromis mossambicus, 466, 470
Ovalipes catharus, 209, 213
trimaculatus, 201
Oxystele variegata, 277
Page 563
Pagurus longicarpus, 278, 282

Palaemonetes pugio, 124, 126, 144
Palinuridae, 486
Pandanus, 407
Panulirus cygnus, 487
longipes, 405, 486, 489, 490, 494
marginatus, 405, 486, 487, 491, 493
ornatus, 405, 486, 487, 489, 490, 494, 496
pascuensis, 405, 486, 492
penicillatus, 405, 486, 487, 489, 490, 492, 493, 494, 496
versicolor, 405, 486, 489, 490, 494
Paphies, 207, 216, 221
donacina, 163, 164, 165, 168, 207, 209, 213, 214, 215, 216, 217
subtriangulata, 163, 164, 165, 168, 207, 209, 213, 214, 215, 216
ventricosa, 163, 164, 165, 168, 208, 209, 210, 211, 212, 213, 214, 215, 216, 218
Paracaesio, 437
Paracentrotus lividus, 145
Paradexamine, 148
churinga, 122, 125, 142, 143
Paradichthyinae, 435
Paraexocoetus, 455
brachypterus, 455
Paralichthys brasiliensis, 188
Paraweldeckia, 137
Parribaccus, 486
antarcticus, 486
caledonensis, 486
holthuisi, 486
Pedunculata, 304
Pelona, 456
Pelvetia fastigata, 148
Penaeus canaliculatus, 405, 495
duorarum, 126
indicus, 490
japonicus, 490
latisulcatus, 491
merguiensis, 405, 490, 495
monodon, 405, 490, 491
semisulcatus, 405, 488, 490, 491, 496
stylirostris, 497
Penicillus capitatus, 148
Periglypta purpurea, 481
Petalonia, 144
Peyssonelia, 269
gunniana, 269
Phaeodactylum tricornutum, 96
Phaeophyta, 134
Phychodrys rubens, 289
Phyllophora pseudoceranoides, 290
Phyllospadix, 110, 111, 133
torreyi, 114
Pinctada margaritifera, 404, 479, 480, 481, 485, 486
maxima, 479
Pinnixia, 167, 172
Pisaster ochraceous, 379
Placuna placenta, 480
Platacidae, 417
Platycephalidae, 469
Platymonas, 372
Plectropomus, 447, 449
leopardus, 414, 425
Plotosidae, 469
Pneophyllum fosliella, 134
Polinices lewisi, 167
Pollicipedinae, 304
Pollicipes, 303–94; 303, 304, 305, 308, 309, 310, 311, 312, 313, 317, 318, 319, 320, 321, 322, 323, 325, 326, 329, 334, 338, 339, 342, 343, 345, 347, 348, 349, 352, 353, 354, 356, 357, 358, 361, 365, 366,
367, 368, 369, 370, 373, 374, 375, 376, 379, 380, 381, 382, 383, 384
cornucopia, 304, 365
elegans, 305, 308, 381; 303, 304, 305, 307, 308, 311, 312, 313, 314, 315, 316, 321, 323, 373, 381
mitella, 304, 329, 361, 369, 372
pollicipes, 307–8, 310, 382–3; 303, 304, 305, 307, 308, 310, 311, 312, 313, 314, 315, 316, 317, 318,319, 320, 321, 322, 323, 326, 329, 332, 333, 334, 339, 342, 343, 350, 351, 352, 354, 355, 356, 357,
359, 360, 361, 362, 365, 366, 367, 368, 370, 372, 373, 374, 375, 376, 378, 379, 381, 382, 383
pollicipes var. minor, 308
polymerus, 306–7, 308–10, 381–2; 303, 304, 305, 306, 307, 308, 309, 311, 312, 313, 314, 315, 316, 317, 319, 320, 321, 322, 323, 324, 325, 326, 327, 328, 329, 330, 331, 332, 333, 334, 335, 336, 337,
338, 339, 340, 341, 342, 343, 344, 345, 346, 348, 349, 350, 353, 354, 355, 356, 357, 358, 359, 360, 361, 362, 364, 365, 366, 367, 368, 369, 370, 371, 372, 373, 374, 375, 376, 377, 379, 380, 381, 382, 383
polymerus: forma echinata, 306, 312
polymerus: forma typica, 306, 312
polymerus madrasensis, 307
sertus, 304
spinosus, 304, 323, 329, 334, 349, 361, 372
Polychaeta, 116
Polydactylus virginius, 181
Polymesoda coaxans, 481
Polynemidae, 404, 467, 468
Polysiphonia harveyi, 143
Pomacanthidae, 431
Pomacentridae, 431
Pomadasyidae, 467, 468
Porifera, 116
Porphyra, 143, 378
Portunus pelagicus, 404
Posidonia, 111, 115, 118, 122, 130, 136, 137, 139, 142
Posidoniaceae, 113
Page 564
Posidonia angustifolia, 113, 114, 137

australis, 111, 113, 114, 115, 118, 119, 133, 134, 136, 137, 139, 140
oceanica, 110, 113, 114, 115, 123, 127, 128, 133, 134, 135, 137, 139, 142, 146, 150, 151
sinuosa, 113, 114, 115, 130, 131, 134, 143, 145
Potamogetonaceae, 113
Pranesus, 456
pinguis, 455
Priacanthidae, 420
Pristipomoides, 432, 437, 439, 440
amoenus, 436
auricilla, 436, 437, 441, 442
filamentosus, 433, 435, 436, 437, 439, 440, 441, 442, 443
flavipinnis, 435, 436, 439, 441, 442
kuskarii, 436
multidens, 436, 437, 438, 439
zonatus, 436, 437, 441, 442
Processa, 439
Pronichthys, 455
Prorocentrum mariallabouriae, 76
micans, 372
Prothalotia lehmani, 121, 128, 137, 138, 140, 142
Prunum apinicum, 127
prunum, 181, 184
Psammobatis, 188
Pteria penguin, 480
Pterocaesio, 456
Punctaria, 143
Pyrene, 128
bidentata, 121, 128, 130, 131, 137, 139, 140, 142, 145
Quindinipagus palatum, 481
Raja, 188
Rastrelliger, 415, 452, 456, 461
brachysoma, 453
kanagurta, 453, 460
Rhabdamia, 456
cypselurus, 460
gracilis, 456, 460
Rhizophora, 400
stylosa, 401
Rhodochorton purpureum, 288
Rhodomela larix, 285
Rhodomelaceae, 133
Rhodophyta, 117, 134
Rhodymenia, 116
Rickettsia, 196
Rinchops nigra, 188
Ruppia maritima, 113, 124, 126
Ruvettus pretiosus, 432, 438
Salmacis sphaeroides, 120, 129, 145

Saloptia powelli, 436
Sardinella, 456, 461
albela, 455
Sardinella marquesensis, 455
Sargassum, 129, 143, 144
muticum, 111, 143, 290
Sargocentron microstoma, 425
Sauridae, 468
Scalpellidae, 304
Scalpelloidea, 304
Scalpellum, 305
scalpellum, 368
Scaridae, 404, 413, 415, 416, 417, 419, 421, 426, 469
Scarus frenatus, 424, 425
globiceps, 424, 425
niger, 424
oviceps, 424
psittacus, 424
rivulatus, 424
schlegeli, 424, 425
sordidus, 424, 425
Scatophagidae, 469
Sciaenidae, 404, 468
Scomberoides, 415
Scomberomorus commerson, 415, 444, 449
Scombridae, 404, 408, 415, 416, 417, 418, 419, 420, 435, 448
Scutarcopagia scobotinata, 481
Scutengraulis hamiltoni, 460
Scylla serrata, 404, 487, 494
Scyllaridae, 405, 486
Scyllarides haani, 486, 491
squammosus, 486, 491, 493
Scytosiphon, 144
lomentaria, 270
Selar, 419, 453, 456, 461
boops, 415, 452, 453
crumenophthalmus, 415, 452, 453, 455, 456, 460
Selenopsis, 181
Semibalanus balanoides, 350, 367, 373, 374, 375, 376
cariosus, 308, 309, 317
Sepiateuthis lessoniana, 481
Seriola dumerilii, 451
Serranidae, 404, 413, 416, 417, 418, 419, 426, 431, 432, 435, 437, 439, 448, 469
Sessilia, 304
Setipinna godavari, 455
Siganidae, 401, 416, 417, 419, 420, 421, 469
Siganus argenteus, 424
canaliculatus, 415, 424
lineatus, 415, 424
rostratus, 424
spinus, 424
vermiculatus, 424
Siliganidae, 469
Siliqua, 164
patula, 163, 164, 165, 168, 172, 173, 174, 175, 176, 177, 178, 179, 217, 218, 219, 220, 221
Page 565
Sinezona, 139
Siphonaria, 269
Skeletonema costatum, 187
Solenacea, 165, 199
Solenidae, 165
Sparidae, 417, 469
Spartina, 278, 280, 282, 283
alterniflora, 278, 280
Sphaeroma rugicauda, 127, 144
Sphoeroides testudinalis, 181
Sphyraena barracuda, 449, 450
queni, 449, 450
Sphyraenidae, 416, 417, 435, 448, 449, 468
Spisula, 481
solidissima, 219
Spondylus nicobarius, 481
Spongomorpha, 144
Spratelloides, 408, 456, 462
delicatulus, 455, 456, 460, 462
gracilis, 452, 456, 458, 460, 462, 503
lewisi, 452, 460
Squilla, 405
Sthenelais boa
Stichopus, 404
chloronutus, 474, 476, 477
horrens, 473
variegatus, 473, 474, 475, 477
Stolephorus, 408, 454, 456, 461
carpenteriae, 460
indicus, 460
insularis, 460
nelsonii, 460
Stomatopoda, 489
Stramonita haemostoma, 246
Strombus canarium, 480
gibberulus, 481
gibberulus gibbosus, 480
lentiginosus, 481
luhuanus, 480, 481
mutabilis, 128, 145
urceus, 480
Strongylocentrotus droebachiensis, 279
pileolus, 473
Synanceia horrida, 431
verrucosa, 431
Synedra, 142
Synodontidae, 469
Syringodium, 111, 137, 139
filiforme, 112, 114, 124, 136, 143, 148
Tachysuridae, 469
Tectarius grandinatus, 481, 182, 483
Tectus pyramis, 481
Telescopium, 481
Tellinacea, 165
Temnopleurus michaelseni, 121
Tethygeneia, 137, 138, 148
nalgo, 125, 142, 143
Tetradita squamosa rubes cens, 309
squamosa rufotincta, 352
Tetraodontidae, 467, 469
Thais emarginata, 241
Thalassia, 111, 115, 118, 137, 139, 141, 142, 151
hemprichii, 113, 114, 120, 129, 133, 134, 145, 150
testudinum, 113, 114, 118, 124, 126, 133, 136, 143, 145, 148
Thalassina anomala, 405, 489
Thalassionema, 85
nitzschoides, 187
Thalassiosira fluviatilis, 86
Thalassodendron, 111, 115, 120
pachyrhizum, 112, 114
Thalotia conica, 121, 128, 130, 137, 139, 140, 142
Thebe pisana, 281
Thelenota, 404
ananas, 474, 475, 476, 477
anax, 474, 477
Theoduxus fluviatilis, 127, 144
Theraponidae, 404, 416, 467, 468
Thoracica, 304
Thryssa, 456, 461
balaema, 460
setirostris, 455, 460
Thunnus alalunga, 450
albacares, 397, 433, 447, 449
obesus, 397, 433, 450
tonggol, 451
Tivelamactroides, 163, 164, 165, 168, 180, 181, 182, 183, 221
stultorum, 163, 164, 165, 167, 168, 169, 170, 171, 174, 217, 218, 219, 221
Toxopneustes pileolus, 473
Tozeuma carolinensis, 126
Trachinotus carolinus, 181, 184
goodei, 184
Trichiuridae, 467, 468
Tricolia, 139
Tridacna, 481
crocea, 480, 481, 482, 483, 484
derasa, 483, 484
gigas, 483, 484
maxima, 481,482, 483, 484
squamosa, 480, 481, 483, 484
Tridacnidae, 480, 481
Tripneustes gratilla, 120, 129, 145, 473
Trochus niloticus, 404, 479, 480, 481
Tryphosella orana, 137
Turbo argyrostomus, 481
chrysostomus, 481
crassus, 480, 481
marmoratus, 404, 479, 480
setosus, 480, 481
Ulva, 116, 143, 144, 145, 146, 149, 269, 271, 284, 287
lactuca, 150, 274
Umbrina coroides, 181, 184
Page 566
Urosalpinx cinerea, 278
Variola louti, 414, 436, 437

Vasum turbinellus, 481
Veneracea, 165
Veneridae, 165, 199
Watssia mossambica, 346, 437, 438
Zebrasoma scopas, 423

veliferum, 423
Zostera, 115, 130, 140, 141, 142, 149, 251, 277, 283
capricorni, 111, 114, 136, 140, 149
marina, 114, 115, 118, 119, 122, 124, 125, 126, 127, 129, 130, 135, 136, 139, 140, 141, 144, 146, 147, 148, 149, 150, 151, 283
mucronata, 114
muelleri, 114
subgenus Zostera, 111
subgenus Zosterella, 111
Zosteraceae, 114
Page 567
SUBJECT INDEX
Oceanography and Marine Biology: and Annual Review 1996, 34, 567–576
UCL Press
References to complete articles are given in bold type; references to sections of articles are given in italics; references to pages are given in normal type.
Advection, 72, 106

Africa,
Atlantic coast, Pollicipes, 308, 310
beach clams, 165, 199
Aleutian Archipelago, Pollicipes, 307
Algae, chemical defences, polyphenols, 286, 287
Allozyme differentiation in littorinids, 239
Allozyme heterozygosity, 243
Allozymes and genetic differentiation, 245, 246
America,
littorinids, 236, 248, 269, 274, 275, 278, 282
Pollicipes, 303
Andaman Sea, physical oceanography, 1–70
Antarctic Bottom Water, 53
Arabian Sea, 2, 3, 10, 22, 33, 39, 41, 43, 50, 52
Arabian Sea High Salinity Water (ASHSW), 33, 43, 50, 51, 52
Arabian Sea Intermediate Water, 52
Archaeogastropods, 237
Arctic packice, littorinids, 271
Articulated calcareous algae, 116
Asia, beach clams, 165, 204
Atlantic,
seagrasses, 114
plankton crossing, 271
Atolls, 399, 400
Australia,
littorinids, 268, 292
seagrasses, 110, 111, 112, 113, 114, 115
AVHRR, 24, 27
Azores, Pollicipes, 308
Balanids,
haemocytes, 329
neurosecretary cells, 326, 327
Barramundi, 464, 465, 471, 472
Bay of Bengal,
climatic classification, 4
physical oceanography of, 1–70
Pollicipes, 307
wind speeds, 5, 6
Bay of Bengal and Andaman Sea,
atmospheric and land environments, 4–10
general climatic aspects, 4–7
net estimates at surface, 9–10
runoff of major rivers, 7–8
coastal oceanography, 10–22
density fields, 19
salinity fields, 13–19
temperature fields, 11–13
turbidity and light, 22
up welling, 19–22
currents and water transports, 56–64
currents, 56–62
transport across different sections, 62–4
deep bay oceanography, 22–56
density fields and mixed layer, 44–50
salinity fields, 35–44
temperature fields, 24–35
turbulent mixing and vertical advection, 53–6
water masses, 50–53
evaporation and precipitation in the bay, 8–10
Bay of Bengal Low Salinity Water, 51
Bay of Fundy, littorinids, 290
Beach clam fisheries, 163–232
Africa, 199–204
white sand mussel, Donax serra, 199–204
biology and ecological role, 200–202
distribution, 199–200
fishery, 203–4
population dynamics, 202–3
Asia, 204–6
wedge clams, Donax cuneatus and Donax faba, 204–6
biology and ecological role, 205
fishery, 206
Australia, 206–7
pipi, Goolwa cockle, Donax deltoides, 206–7
distribution, 206
fishery, 207
population dynamics, 207
Caribbean, 180–56
beach clams, chipichipi, Donax denticulatus and D.striatus, 182–6
fishery, 186
Page 568
guacuco, Tivela mactroides, 180–82

distribution, 180
fishery, 182
Discussion, 217–21
future prospects, 221
future research needs, 219–27
densitydependent processes, 220
disturbance associated with harvesting and
incidental mortality, 220
modelling and management, 220
recruitment processes, 219–20
general, 217–79
Europe, 194–9
wedge clam, Donax trunculus, 194–9
fishery, 198–9
key features tabulated, 168
New Zealand, 207–16
toheroa, Paphies ventricosa, 208–13
distribution, 208
fishery, 211–13
tuatua, Paphies donacina, 215–16
distribution, 215
fishery, 216
tuatua, Paphies subtriangulata, 213–15
distribution, 213
fishery, 214–15
North America, 165–80
Pacific razor clam, Siliqua patula, 172–80
distribution, 172
fishery, 175–80
Pismo clam, Tivela stultorum, 165–71
distribution, 165
fishery, 169–77
southern South America, 186–94
macha, Mesodesma donacium, 191–4
fishery, 193–4
yellow clam, Mesodesma mactroides, 186–91
fishery, 190–91
Beach clams,
as substratum,
for algae, 184
for gastropod eggs, 181
for hydroids, 167, 181, 184
birds as predators on, 167, 173, 184, 188, 201, 205, 207, 209, 213, 215, 218
crabs as predators on, 167, 173, 181, 184, 197, 201, 205, 209, 213, 218
disturbance by motor vehicles, 168
fishes as predators on, 167, 173, 181, 184, 188, 197, 201, 207, 209, 213, 218
gastropods as predators on, 167, 181, 184, 188, 197
geographical distribution, 164
isopods as predators on, 188
management of fisheries, 169, 170, 171, 175, 176, 177, 178, 180, 186, 190, 191, 193, 199, 203, 204, 207, 212, 213, 215, 216, 219, 220, 221
migration, 183, 187, 196, 200, 204, 206, 209, 213, 215, 217
occurrence in shell middens, 169, 175, 182, 203, 207, 211, 214
pigs as predators on, 184
polychaetes as predators on, 181, 207
sea otters as predators on, 167, 168, 173
taxonomy and common names, 165
Bêchedemer, 473, 474, 475, 476, 478, 479, 502, 505, 507
Belgium, littorinids, 245, 249
Bering Strait,
littorinids, 235
Pollicipes, 302
Biology of the Littorinidae,
evolutionary aspects, 233–62
role in ecology of intertidal and shallow marine ecosystems, 263–302
Birds,
as predators of littorinids, 238
as predators on beach clams, 167, 173, 184, 188, 201, 205, 207, 209, 213, 215, 218
Blennies, as predators of littorinids, 238
Bodega Marine Reserve, California, 379
Brahmaputra River, 2, 4, 7, 13, 16, 18, 19, 35, 50, 61, 64
Britain,
littorinids, 245, 247, 272, 287, 289
Pollicipes, old records, 307
California,
littorinids, 276
Pollicipes, 305, 306
Canada,
littorinids, 271, 286, 287
Pollicipes, 359, 361, 371, 379, 381, 382
Canary Islands, Pollicipes, 308
Caribbean,
Page 569
littorinids, 244
seagrasses, 112, 113
Cauvery River, 2, 4, 7
Cement, barnacle
properties of, 321
solidification of, 322
Cement discharge, Lepas, 321
Cestodes, effect on sexual development of Pismo clam, 167
ChilakaLake, 11, 12, 14, 19
Chile,
seagrasses, 114
no Pollicipes, 305
China, Pollicipes, 308
Chipichipi, Donax denticulatus and D.striatus,
feeding, 184
fishery, 186
growth rates, 184, 185
larval development, 184
predators on, 184
recruitment, 185
reproduction, 183
vertical distribution, 182, 183
Cirripedes,
hatching substance, 378
utilization of lipids, 352, 353
Coastal fisheries, Pacific Islands, 395–531
Colour polymorphism, littorinids, 237, 238
Coral reefs, S.Pacific, 400
Corticated macrophytes, 116
Crabs,
as predators of littorinids, 238, 240, 241, 242, 246
as predators on beach clams, 167, 173, 181, 184, 197, 201, 205, 209, 213, 218
Crustaceans,
as grazers on seagrasses, 119, 120, 122, 123
Damodar River, 2, 4, 7
De Lury’s method, 427
Denmark, littorinids, 247, 289
Department of Fisheries and Oceans, permits for harvesting P.polymerus, 381
Detritus, defined, 116
Diel variation,
in seagrass epifaunal grazers, 138, 139
Diffusional boundary layer, 73, 74
Domoic acid, in beach clams, 178
Drift algae, 116
Eastern Dilute Waters of IndoPacific Origin, 50

Echinoderms, as grazers on seagrasses, 120, 121
ECOPATH model, 430
Ekman layer, 21, 55, 56
Ekman transport, 21
El Niño Southern Oscillation (ENSO), 402, 403
effect on beach clams, 170, 177, 192, 193
Electrophoretic techniques, in genetic differentiation, 236
ELEFAN, 192
Encrusting algae, 116
Endemism, in seagrasses, 111
England, littorinids, 276, 277, 278
Ensenada, Lower California, Pollicipes, 306
Enzyme polymorphism, 243
Epiphytes,
beneficial effect on seagrasses, 119
defined, 116
detrimental effects on seagrasses, 118
Europe,
littorinids, 235, 236, 242, 247, 265, 269, 271, 272, 273, 279
Pollicipes, 303, 307, 339, 361, 362, 365, 366, 367, 378, 382, 383
Eutrophic conditions, impact on seagrass grazerepiphyte interactions, 150–51
Exclusive Economic Zone (EEZ), 397, 433, 434, 438
FAO, 452, 453

Filamentous algae, 116
Filterfeeding molluscs, in seagrass beds, 121, 122
Fish aggregating devices (FAD), 444, 446, 447, 452
Fisheries, Pacific islands, 395–531
crustaceans, 486–97
echinoderms, 472–8
finfishes,
deepslope, 432–44
estuarine, 463–72
nearshore, 444–63
reef, 408–32
molluscs, 478–86
Fisheries research, Pacific islands, 500, 501, 502, 503, 508
Fishes,
as grazers on seagrasses, 119, 120
as predators on beach clams, 167, 173, 181, 184, 188, 197, 201, 207, 209, 213, 218
Fishing methods, Pacific islands, 405–8
Foliose algae, 116
Fox’s production model, 429, 430, 441, 461
France,
Pollicipes, 307, 310, 350
Ganges (Ganga) River, 2, 3, 4, 7, 10, 11, 13, 16, 18, 19, 21, 35, 50, 57, 61, 64
Germany, littorinids, 292
Giant clams, 480, 482, 483, 484, 502, 503
Godavara River, 2, 4, 7, 10, 11, 19, 35, 56, 57, 58, 61, 62, 64
Golgi apparatus, 367, 377
Goolwa cockle, See Pipi,
Goose (or gooseneck) barnacle, origin of name, 303
Grazerepiphyte interactions in temperate seagrass meadows, factors affecting, 109–62
Page 570
conclusions, 151–52
general biology, 110–31
epiphytes, 116–19
interactions between epiphytes and seagrasses, 117–19
techniques for measuring abundance, 117
grazers, 119–31
food selection and feeding patterns, 122–3
measurements of diets, 123
types of, 120–21
seagrasses, 110–15
grazerepiphyte interactions, 141–51
effect of epiphytes on grazers, 147–9
food, 147
habitat, 147–9
shelter from predation, 149
effect of grazerepiphyte interactions on seagrasses, 149–57
impact of eutrophic conditions on grazerepiphyte interactions, 150–51
effect of grazers on epiphytes, 141–7
change in species composition, 146
filter feeders, 144–7
grazing and epiphyte production, 142–6
removal of epiphyte biomass, 141–2
spatial and temporal patterns of abundance of epiphytes and their grazers, 132–41
epiphytes, 132–5
spatial patterns, 132–4
temporal patterns, 134–5
grazers, 135–41
Grazers,
within seagrass meadows, diversity, 119
of seagrass epiphytes, studies of, tabulated, 124–31
Great Barrier Reef (GBR), 414, 423, 424, 425, 426, 431
Great Channel, 2
Greenland, littorinids, 271
Growth rate index ф′, for beach clams, 168, 169, 173, 189, 192, 197, 202, 207, 214
Guacuco, Tivela mactroides,
as substratum for gastropod egg capsules, 181
feeding, 181
fishery, 182
growth rate, 181, 182
mortality, 182
predation on, 181
reproduction, 180, 181
vertical distribution, 180
Gulf of Aden, 3
Gulf of Mannar, 2, 10
Gulf of Maraban, 57
Gulf of Mexico,
seagrasses, 112
Habitat complexity,
of seagrass epiphytes, 148, 149
HardyWeinburg equilibrium, 243, 244
HardyWeinburg rule, 243
Hawaii, littorinids, 242
Helgoland, littorinids, 271
Heterozygosity, littorinids, 254
Hong Kong, littorinids, 269, 292
Hydroids, commensal on beach clams, 167, 181, 184, 215
Ibla,
muscles in, 322
nervous system of, 323
Iceland, littorinids, 271
Indian Equatorial Intermediate Water, 4, 50
Indian Monsoon Current, 18, 23, 35, 57, 61
Indian Ocean, 1, 2, 3, 10, 22, 23, 34, 53, 55, 57, 60
seagrasses, 112, 113
Indian Ocean Bottom Water, 52
Indian Ocean Equatorial Water, 50
Infectious gill disease (NIX), in Pacific Razor Clam, 172, 174, 176
Intermediate High Salinity Water (IHSW), 5, 52
International Indian Ocean Expedition (IIOE), 1, 3, 8, 23, 50, 61, 64
Intertidal pedunculate cirripedes, 303
Irrawady River, 2, 3, 4, 5, 7, 10, 11, 17, 19, 36, 43, 44, 61, 64
Isopods, as predators on beach clams, 188
Jan Mayen, Pollicipes, 308
Kolmogorov length scale, 92, 93, 98, 100, 105

Krishna River, 2, 4, 7, 10, 11, 19, 35, 44, 61, 62, 64
Lecithotrophic development, 252

Lecithotrophic larvae, 373
Lepadids, circulatory structures, 327
Lepas, composition of shell plates, 353
Leslie’s stock depletion model, 427, 441, 442
Lithotrya,
blood pump, 328
muscles in, 322
Littorina littorea, densities, 273, 274
Littorinidae,
community interactions, 265–90
defining species, 235–7
detritivores, 280–83
Littoraria irrorata, 280–83
climbing behaviour, 280–81
feeding, 282–3
predation, 281–2
direct developing species, 239–41
Littorina saxatilis, 239–40
Littorina sp., 241
epilithic grazers, 266–79
high shore, Nodilittorina unifasciata, 267–9
low shore, Littorina littorea, 271–9
Page 571
community effects, 272

effects on algae, 272–6
effects on nongrazing motile animals, 278
effects on other grazers, 276–8
habitat modifications, 278–9
historical background in north America, 271–2
mid shore, Bembicium nanum, 269–71
epiphytic grazers, 283–5
Littorina mariae, 284–5
evolutionary aspects, conclusions, 252–4
grazing, 265
heterozygosity, 243–8
evidence against theory, 247–8
evidence in support of theory, 245–6
theory, 243–5
larval development, 242–52
dispersal, 248–52
life history theory, 250–52
macroalgal grazers, 285–90
algal defences against grazing, 285–6
Lacuna spp., 289–90
Littorina obtusata, 287–8
morphological variability, 237–42
phylogeny and ecology, 264–5
planktonic developing species,
Littorina littorea, 241–2
Nodilittorina hawaiiensis, 242
role in the ecology of intertidal and shallow marine ecosystems,
conclusions, 292
summary, 291
shell shape, 239
taxonomy and evolution, 234–5
Littorinids,
barnacle morphs, 239
algal sporelings, 288
escapes, 266, 267, 270, 271, 274, 275, 285
allozyme studies, 230
and mucous trails, 272
Australia, 235
barnacles inhibit grazers, 264
chemical defences, 263
climbing as predator avoidance, 281
colour polymorphisms, 233, 234
crab predation, 281, 287
densities, 277, 290
detritivores, 266, 263
diets, 265
effect of barnacles, 268
effect of Zostera beds, 277
epilithic grazers, 263, 264, 266
epiphytic grazers, 263, 266
gut contents, 282, 283
heterozygosity, 233, 236
in Holocene period, 238
introduced by shipping, 245
larval transport across Atlantic, 248, 249
macroalgal grazers, 266
major feeding types, 266
morphology, selective processes, 237
mucous trails, 280
northern temperate region, 235
on mangroves, 235, 264, 280
ovovivipary, 249, 252
pollutants, 264
predation,
by conchs, 281, 282
by crabs, 267, 269, 284
by fish, 284
predation exclusion experiments, 281
predators of, 238
preying on egg masses, 277
rk selection and bethedging, 250, 251
shells used by hermit crabs, 278, 282, 283
shelter for young provided by barnacles, 276
southern temperate origin, 235
taenioglossan radula, 265
taxonomy, cladistic approach, 234
use as sentinel organisms for testing pollutants, 264
Macha, Mesodesma donacium,

artisanal fishery, 193
growth rate, 192
recruitment, 192
reproduction, 192
vertical and longshore distribution, 191
Macroalgae, 116
Mahanadi Water, 2, 4, 7, 16, 18, 19, 21, 56, 57, 58, 61, 64
Mahaweli River, 4, 7
Malacca Strait, 1
Mangroves, S.Pacific, 400, 401
Marine Station of University of California, Santa Barbara, 306
Mediterranean,
seagrasses 113, 115
Pollicipes, 308, 310
Meghna River, 7
Mexico, Pollicipes, 305, 308
Microalgae, 116
Molluscs,
Mortensen Expedition, 306
Motherof pearl, 478, 479, 480, 485, 505, 507
Nematodes, in beach clams, 196

Nemerteans, commensal in Pacific razor clam, 172
New England, littorinids, 239, 246, 264, 270, 271, 272, 275
New South Wales, littorinids, 267, 270, 271, 276
New Zealand,
littorinids, 268
Pollicipes, 361
seagrasses, 114
Page 572
Ninety Degree East Ridge, 2

NOAA, 27
Nonlethal (grazing) predation, 181, 184, 197, 201, 213
North America,
littorinids, 234, 235, 246, 247, 271, 272, 273, 292
Pollicipes, 338, 350, 357, 358, 359, 361, 362, 363, 366, 367, 373, 379, 380, 382, 383
seagrasses, 114
North Atlantic, littorinids, 234, 235, 245, 246, 249
North Equatorial Current, 2, 3, 18, 21, 32, 38, 41, 43, 47, 48, 57, 58, 59, 61
North Indian Ocean Deep Water, 50, 52
North Pacific, Pollicipes, 307
Northern Dilute Water, 50
Norway, littorinids, 273, 279
Nova Scotia, littorinids, 275, 283, 288
NOVARA, 3
Nuclear inclusion X (NIX), 173, 174, 176
Nusselt number, 77, 79
Nutrient fluxes to planktonic osmotrophs,
in the presence of fluid motion, 71–107
mass transfer to cells in stagnant water, 73–86
motile cells: swimming and sinking, 75–86
application of theory to swimming and sinking
cells, 79–84
effect of shape on the nutrient flux, 84–5
theory, 75–9
nonmotile cells: pure diffusion, 73–5
nutrient uptake by motile cells in stagnant water,
observations, 85–6
mass transfer to cells in steady shear flow, 86–91
laboratory experiments, 90–91
theory, 86–90
nonmotile cells in steady shear flow, 86–90
valuation of the theory in an engineering
context, 90–91
mass transfer to cells in turbulent water, 91–9
application to the ocean, 95–6
nonmotile cells in turbulent flow, 91–5
translational motion in turbulent water, 96–9
flowinduced translational motion, 98–9
swimming or sinking in turbulent water, 96–7
ORSTOM, 429
ORV SAGAR KANYA, 23
Otoliths, 422, 423, 424, 425, 440, 503, 508
Ovoviviparity, in littorinids, 242
Pacific islands, coastal fisheries, 395–531

Pacific razor clam, Siliqua patula,
burrowing 172
commercial fishery 175, 176, 177, 178
feeding 173
in shell middens, 175
longevity, 173
mortality, 174
predation on, 173
recreational fishery, 176, 177, 178
recruitment, 173
reproduction, 172
Pacific,
littorinids, 235, 241, 246
seagrasses, 112, 113, 114
Paddle weed, 115
Palar River, 7
Palk Bay, 10
Paralytic shellfish poisoning (PSP), and beach clams, 171, 175, 178, 204, 219
Pea crabs, commensal in beach clams, 167, 172
Pearls, 478, 479, 480, 486
Peclet number, 77, 78, 79, 81, 82, 85, 86, 89, 94, 105
Peclet numbers, for phytoplankton, 76
Pedunculate cirripedes of the genus Pollicipes, 303–94
Pennar River, 2, 4, 7
Periphyton, defined, 116
Persian Gulf, 3
Persian Gulf Water (PGW), 51, 52
Peru, Pollicipes, 305
Phenolics,
release by seagrasses inhibits epiphyte growth, 118
Photoreceptors in balanids, 322
Physical oceanography, Bay of Bengal and Andaman Sea, 1–70
Pigs, as predators on beach clams, 184
Pipi (Goolwa cockle), Donax deltoides,
fishery, 207
growth rate, 207
predation on, 207
reproduction, 206
Pismo beach, 168, 169, 170, 171
Pismo clam, Tivela stultorum,
burrowing, 167
commercial fishery, 169
fecundity, 166
feeding, 167
in middens of coastal Chumash Indians, 169
longevity, 168
mortality, 169, 171
predators on, 167
recreational fishery, 170, 171
recruitment, 168, 171
Planktotrophic larvae, 373
Page 573

Point Conception, nearshore currents, 363, 364
Pollicipes,
growth zone, 354
rosette pattern as protection from predators, 380
abnormal shell form, 317
action of mouthparts, 342
adhesion and cement, 321–2
allocation of energy, 345–7
and mussel communities, 308, 309, 310
arrangement of capitular plates, 312
biochemical composition of embryos during development, 352
capitular plates, 317, 318, 319
cement ducts, 321
chemical composition, 350–53
of bodies and egg lamellae, 350–53
of calcified structures, 353
chromosomes, 369
circulation of haemolymph during contraction and elongation of peduncle, 340
circulatory system, 327–34
cirral activity and feeding, 339–47
classification of, 303, 304
commercial exploitation, 381–3
P.elegans, 381
P.pollicipes, 382–3
P.polymerus, 381–2
composition of shell plates, 353
copulation, 369–70
description of adults, 311–17
description of three species, tabulated, 313, 314, 315, 316
details of breeding seasons, 366
details of reproductive periods, 361
development of capitular plates, 319, 320
development of peduncular scales, 354, 355
ecological habitat, 308–9
effect of pollution, 380–81
effect of temperature change and desiccation, 347–8
effect of water currents on movement of peduncle, 335, 336, 337, 338
embryo development, 370–72
endocrinology, 326–7
endolithic algae, 378
energy values during embryo development, 352
fatty acid composition of lipids, 351
fertilization, 369–76
food items found in midgut, 344
fracturing of lamina, 317
geographical distribution, 305–8
growth rates, 359, 360, 361
haemolymph, 329–31
pressure, 331–3; 322
properties of, 329, 330, 331
haemolymph pump, 328, 329
in Upper Cretaceous sediments, 304, 305
ingestion of Artemia, 345, 346
inner integument, 320–21
integument and calcified parts, 317–21
larvae, use in marine bioassays, 334
lipids, trigyceride and phospholipid composition, 350
mechanical stimulation of cirri, 340
moulting, 320
moulting and growth, 353–61
mouthparts and digestive system, 342–3
movement of juveniles on adult peduncle, 357
muscles, 322
naupliar development, 372–3
nervous system, 322–7
neurosecretory cells, 326
number of broods per year, 366
number of eggs per brood, 366
ocellar nerves, 324, 325, 326
oogenesis, 369
orientation and movement of peduncle, 334–9
outer integument, 317–18
adults, 318–19
juveniles, 319–20
oviducal sac, 373–6
composition of, 376
ovisacase, properties of, 375
oxygen uptake of egg lamellae, 349, 350
pedicel and penis, 367, 368
peduncular scales, 317, 318
peristaltic constrictions during early embryonic
development, 377
photoreceptors, 325–6
physiological races, 363, 364
potential predators, 379, 380
predation, 379–80
rate of cirral beating in juveniles, 342
rate of water loss from peduncle, 348
recolonization of denuded areas, 358
rehydration, 348
relative distribution of mono and polyunsaturated acids in lipids, 352
respiration, gaseous exchange and oxygen uptake, 348–50
rhythmic activity of stalk nerves, 325
rigidity of peduncle, 333
separation of three species by Cenozoic movements of the continents, 305
settlement of cyprids on peduncle, 356, 357
size at maturity, number of broods and number of eggs, 361–6
size of young, 366
sizes of nauplii and cyprids compared, 373
spermatogenesis and spermatozoa, 366–8
stages of embryo development compared, 371
stomach contents, 344–5
strength of peduncle, 333–4
structure of spermatozoa, 367
transference of food from cirri to mouth parts, 341, 342
Page 574
ultrastructural investigations, 376–8

use of radioisotopes to study growth, 355
use of silicon rubber to elucidate circulatory system, 327
Pollicipes elegans,
ecological habitat, 308
fishery for in Costa Rica, 381
Pollicipes pollicipes,
consumption rate in Spain, 383
ecological habitat, 310
overfishing, 383
remains found in Neolithic hearths, 382
Pollicipes polymerus,
ecological habitat, 308–10
fisheries for in north America, 381
Polychaetes, as predators on beach clams, 181, 207
Polymorphism, in littorinids, 237
Polyphenols, in algae, 274
Portugal, Pollicipes, 308, 310
Predation, and littorinids, 238, 252, 253
Preparis Channel, 2, 43, 61
Puget Sound, Pollicipes, 306
Rainfall,
effect on catches, 462, 295
South Pacific islands, 401, 402
Red Sea, 55
Red Sea Water (RSW), 51, 52
Reproductive strategies, littorinids, 233
Resurgent populations, 168
Reynolds number, 75, 77, 94, 98, 100, 105
Reynolds numbers, for phytoplankton, 76
RMIS INVESTIGATOR, 3
RV GAVESHANI, 3, 23, 64
Sakhalin Island,
littorinids, 238
Pollicipes, 307
Salween River, 2, 3, 4, 5, 7, 19, 61, 64
Santa Barbara oil spill of 1969, 380
Schaeffer’s stock production model, 429, 441, 461
Scotland, littorinids, 284
Scripps Institution of Oceanography, 306, 359
Sea nymph, 115
Sea otters, 167, 168, 173
Sea urchins,
Seagrass grazers,
larger scales, 137–8
latitudinal patterns, 138
small scales, 136–7
diel patterns, 138–9
seasonal patterns, 139–41
Seagrass meadows, temperate, factors affecting grazerepiphyte interactions in, 109–62
Seagrass epiphytes,
rates of grazing on, tabulated, 143, 144, 145
seasonality, 135
larger scales, 134
small scales, 132–4
chronological sequence of development, 133
orientation of the leaf surface, 133–4
relative age of the leaf, 132–3
Seagrasses,
distribution, tabulated, 112–14
ecological characteristics, tabulated, 112–14
leaf characteristics, tabulated, 112–14
morphological characteristics, tabulated, 112–14
morphological classification of, 111, 115
salinity tolerances, tabulated, 112–14
south Pacific, 401
Seamounts, 442, 503
Seasonality,
in seagrass epifauna, 139, 140, 141
in seagrass epiphytes, 135
Sherwood number, 77, 78, 80, 89, 94, 105
Shrimp fisheries, Pacific islands, 488, 490, 492, 493, 495, 496, 497
South Africa,
Donax, 199, 253
South America,
Pollicipes, 305
South Pacific Commission, 397, 406, 432, 434, 435, 450
South Pacific Islands,
climate, marine environment, 401–3
finfish fisheries, 408–72
deepslope fisheries, 432–44, 506
catch composition, 434–8
catch rates, 438–40
description, 432–4
fisheries biology, stock assessment, 440–2
socioeconomic developments, 443–4
estuarine fisheries, 463–72
catch rates, 470
fisheries biology, stock assessment, 471
socioeconomic developments, 472
nearshore fisheries, large pelagic species, 444–52
catch composition, 447
nearshore fisheries, small pelagic species, 452–63
Page 575
socioeconomic developments, 463
reef fisheries, 408–32
fisheries production, 497–500
fishing methods, 405–8
industrial scale, 408
subsistence and artisanal, 405–8
invertebrate fisheries, 472–97
crustaceans, 486–97
echinoderms, 472–8
molluscs, 478–86
map of area, 396, 398
physical geography, 399–401
South Pacific, fisheries resources, 403–5
Southern Bay of Bengal Water, 50
Southern Ocean, littorinids, 249
Spain, Pollicipes, 307, 310
Spiny lobsters, 486,487,489, 490, 491, 492, 493, 494, 496, 497
Spitsbergen, fossil littorinids, 271
Sporozoans, in beach clams, 196
Surf zone blooms, 173, 201, 202
Sweden, littorinids, 245
Temperate seagrass meadows, factors affecting grazerepiphyte interactions in, 109–62

Ten Degree Channel, 2, 5, 45
Tethys Sea, littorinids, 235
Toheroa, Paphies ventricosa,
burrowing, 208, 209
commercial fishery, 211
growth rate, 209
longevity, 209
predation on, 209
recreational fishery, 211, 212, 213
recruitment, 209, 211
reproduction, 208
Tourism in Pacific islands, 505
Trematodes, in beach clams, 167, 184, 196
Trochus, 479, 482, 483, 484, 485, 507
Tuatua, Paphies donacina,
burrowing, 215
feeding, 215
fishery, 216
growth rate, 216
longevity, 216
predation on, 215
recruitment, 216
reproduction, 215
Tuatua, Paphies subtriangulata,
burrowing, 213
feeding, 213
fishery, 214, 215
growth rate, 214
predation on, 213
recruitment, 214
reproduction, 213
Upwelling Index, 20, 21

Upwelling, Bay of Bengal, 10, 11, 19–22, 65
VALDIVIA, 3
Venice, littorinids, 236, 239, 245
Wadden Sea, littorinids, 271

Wales, littorinids, 248
Water Mass E, 51
Wedge clam, Donax trunculus,
burrowing, 196
feeding, 197
fishery, 198, 199
growth rate, 197
longevity, 197, 198
mortality, 197
predation on, 197
Wedge clams, Donax cuneatus and D.faba,
burrowing, 205
fishery, 206
growth rate, 206
predation on, 205
reproduction, 205
Page 576
White sand mussel, Donax serra,

burrowing, 201
commercial fishery, 203, 204
feeding, 200, 201
growth rate, 202
in stone age shell middens, 203
mortality, 203
caused by red tide, 203
predation on, 201
recreational fishery, 190, 191
reproduction, 200
Wire weed, 115
Woods Hole, MA, littorinids, 277
Yellow clam, Mesodesma mactroides,

artisanal fishery, 190, 191
fecundity, 187
feeding, 187, 188
growth rate, 189
mortality, 188, 189
predation on, 188
recruitment, 188
reproduction, 187
vertical and longshore distribution, 186, 187
Zostera meadows, littorinids in, 251

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(Oceanography and Marine Biology - An Annual Review) Alan Ansell - Oceanography and Marine Biology - An Annual Review, Volume 34-CRC Press (1996)

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OCEANOGRAPHY and MARINE BIOLOGY

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OCEANOGRAPHY and MARINE BIOLOGY

The Dunstaffnage Marine Laboratory Oban, Argyll, Scotland

© A.D.Ansell, R.N.Gibson and Margaret Barnes, 1996

ISBN 0­203­50126­8 Master e­book ISBN

ISBN 0­203­23132­5 (OEB Format)

Physical oceanography of the Bay of Bengal and Andaman Sea 1

Author index 533

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This page intentionally left blank.

PHYSICAL OCEANOGRAPHY OF THE BAY OF BENGAL AND

Oceanography and Marine Biology: an Annual Review 1996, 34, 1–70

Atmospheric and land environments

General climatic aspects

Runoffs of major rivers

Annual mean (m3 s−1) Total run off volume (×109m3)

Evaporation and precipitation in the Bay

Latitude belt (°N) Longitudinal belt (°E)

Net estimates at surface

Table 3 Precipitation (cm/season and cm/yr) over the Bay.

Figure 5 Distribustion of temperature during winter at various levels in coastal areas.

Figure 6 Distributions of salinity during winter at various levels in coastal areas.

Figure 7 Distributions of temperature during summer at various levels in coastal areas.

Figure 8 Distributions of salinity during summer at various levels in coastal areas.

Turbidity and light

Deep bay oceanography

100 metre surface

200 metre surface

500 metre surface

1000 metre, 2000 metre and 3000 metre surfaces

100 metre surface

200 metre surface

500 metre surface

1000 metre, 2000 metre and 3000 metre surfaces

Density fields and mixed layer

100 metre surface

500 metre surface

1000 metre, 2000 metre and 3000 metre surfaces

Turbulent mixing and vertical advection

Figure 40 Vertical profile of mean KZ in the northeastern Indian Ocean.

Currents and water transports

Figure 42 Geostrophic current marks transient cells.

Figure 43 Geostrophic current vectors marks transient cells.

Figure 44 Dynamic topographies of different surfaces during summer.

ing summer and winter, respectively.

Transport across different sections

Marine Sciences 5, 179–89.

NUTRIENT FLUXES TO PLANKTONIC OSMOTROPHS IN THE

Oceanography and Marine Biology: an Annual Review 1996, 34, 71–107

Mass transfer to cells in stagnant water

Non­motile cells: pure diffusion

Motile cells: swimming and sinking

Organism Cell radius Swimming speed Sinking speed Re Pe Source

For Pe 1 (Acrivos & Goddard 1965)

Application of the theory to swimming and sinking cells

Predicted cell radius (mm)

Flow generated by Pe<<1 Pe>>1

Effect of shape on the nutrient flux

Observations: nutrient uptake by motile cells in stagnant water

Mass transfer to cells in steady shear flow

Non­motile cells in steady shear flow

ISBN 0203501268 Master ebook ISBN

ISBN 0203231325 (OEB Format)

Nonmotile cells: pure diffusion

Nonmotile cells in steady shear flow

Nonmotile cells in turbulent flow

Flowinduced translational motion

Appendix 2 Advectiondiffusion model

FACTORS AFFECTING GRAZEREPIPHYTE INTERACTIONS IN

Effect of grazerepiphyte interactions on seagrasses

Impact of eutrophic conditions on grazerepiphyte interactions