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Sandy Beach Ecology A Review
Sandy Beach Ecology A Review
Sandy Beach Ecology A Review
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SANDY
BEACH
ECOLOGY
REVIEH
ANTON McLACHLAN (Zoology Department, University of Port Elizabeth, P.O. Box 1600, Port Elizabeth 6000, South Africa)
sand lagoons
flats out
and assets
-;ea.
coastlines
impor~ant
where
they
re?resent
recreational
Nevertheless,
in estuaries CJr closed only open marine LJeaches. sucb beaches may differ in their Jegree of exposure
.:-ocJuctive
..::ommercially. regarc1ed
zones against the buffer they areas are very some exploited are and However, they have been marine and T.1!ere deserts largely
considerably
to wave action. On a 20 point exposure scale, developed studies for on (McLachlan, intertidal fauna 19800.) , scoring 11 to 18 are considered by are characterised and continuous, often heavy, 'v-lave action, the absence of silt, a mobile fauna and a high degree of oxygenation of the sand. (scores Sheltered beaches have 5-10) less,
fa Ulia
as
uy
many
biologists
neglected
beaches exposed
began studies on the Tbe vl0 r k of Pearse also pioneering and first a
sandy
b 1 y,
the
qualitative whole
b,::ach
to
Since
c e d
evaluate the n
sid era
often
wh i c b
intermittent
rnay cons t rue t
i.4aVe
sern i -pe
C3. c
t i
'8
r-~
beach ecology
tho ugh i t has
rrnanen
[-, s a
a 1 v7 a
\; a c1 an
y s
con
GurroVls
and
reduced
layers to
present
and
behind other aspects of coastal sarine ecology in the attention During this period it has enjoyed. research on beaches has biological early from taxonomic spread and
~alitative ol09:l
.species
lagged
the surface. 5 points on this scale are not open ocean beaches
Ge 1 0\-1
fall
beacbes
outside Hright,
this
tOVlards today.
extremes based on morphodynamics. As the sandy beach occupies the a dynamic adjacent
approach
The
term
sandy
beach
has
been
loosely
interface its
in the literature to cover a range '-'sed of environments from high energy open to extremely sheltered ocean beaches
est ua r i ne sand
boundaries
flats. a
of
this
review
terrestrial and marine environments are not always clear and the functional extent of the beach itself bas seldom sand beach discussed. The Dee" considered here to be part a system
considered
to be a sandy littoral
322
comprising highest boundary of the envelope limit they which ecosystem, tropical not sandy interactions landvlard seavlard situations, driven inshore features in Fig. 1. by water. of include and of exist. I drift out waves of
i)
been and
written relationships
seeJimentary
moving the
surf circulation
zone
surf
cells
to The
This call is the are boundary boundary surf breaking The of the
temperate
coasts,
eJate
has
on and
the most
of
intertidal of
as a whole,
intertidal lJeaches substratum provides
open lJy
topography
this
system
are
illustrated
subst
..
DUNES
DUNES ACTIVE DUNES
BEACH
II
I J
NEARSHORE
INNER TURBULENT ZONE
A
'\ (
'STABLE I ~
FORE SHORE
OUTER TURBULENT
ZONE
FIGURE 1. Profile the text. The will other physical not be papers
of
a typical
of
this
system Besides
here. volume,
descript-
consisting
of
the
sand
surface
and upper layer of sediment and ii) the interstitial habitat consisting of the
porous
former, is a
system
un ifo rm
of
lJu t
the
sand
and
lJody.
The
particularly
on exposed
lJeaches,
unstable,
Inman,
dynam ic
323 essentially tyJO dimensional latter is a more stable system. The and complex environment. macrofauna (Hummon these et are Thus although meiofaunainteractions may occur a1. , Reise, 1979) 1976; generally of negligible flow on
habitat with clear vertical gradients and is essentially a three dimensional system (1)1cLachlan,1977a). It has great vertical extent on exposed beaches or high energy 'windows' where strong hydrodynamic deep into the forces introduce oxygen sediment. Elsewhere its layers reduced
The only review on sand beach ecology as a whole is some that of Hedgpeth of veach (1957), ecology Both although aspects
are covered
lJy Eltringbam
(1971).
':2;;e macrofauna of sandy beaches and the (and microfauna) of meiofauna their entirely comprise two interstices components with separate faunal virtually no overlap or exchanges of energy. This is because the meiofauna forms small interstitial are extremely 'while the macrofauna are several orcJers of
mag:iitude
1
these works are outdated and there is a a synthesis distinct need for of information on sandy lJeaches. The aIm of this review is to outline our knowledge of the ecology of individual components structure ecosystems through of sandy beach biotas and the and as the a function whole. It
interstitial
of beach progresses
larger
1971 ;
disting-
~.'1cI:1tyre
Murison, Schwinghamer
McL&chlan,
:977a) .
the surf zone and birds of the beach and dune nlargin to functional aspects within energy the f r ame'v';o r k of flow and nutrient cycling.
2. The THE INTERSTITIAL SYSTEM
meiofauna size,
and
rnacrof
auna.
As of gap
sediments
mean
grain
become finer below 200 urn large lJur roy/ing forms increasingly become meiofauna and
'v"lee n
important
bet
tend
to
bridge
the
size
and meiofauna. macrofauna separation into two makes Thi s components less marked in finer and facilitates better energy sediments
101,0.'
between the sand averages about 40% of Most the total sediment volume. physical properties of this system are grains generally directly c1etermined by the sediment properties which in turn are related to the wave and current regime the geological history of
as vlell
porous
system
Energy between these components. components also between these meiofauna because the are increases surface the and concentrated near
f f 1 01,01
an
as area.
Grain size, shape and sorting are most important in fixing porosity and permeability Drainage which influence drainage. the organic layers. coarser and is critical in determining
macrofauna
abundance
deposit in sediments
increase
in In
finer
~ediments.
coarser
intermedi",te
because they forms are absent, represent a size range selected against by the physical force s of the beach
moisture content, oxygen and input and the depth of reduced Permeability substrate increases and better with
sorting
324 also increases on steeper drainage beaches. The interactions between these and covered factors are well documented in several papers (Hebb, 1958; Hulings, Gray, 1971; Crisp, Hilliams, 1971). The major interstitial process involving the system on open beaches is
the filtration of sea water. ':Chis water the sediment either is introduced into by flushing by waves and tides in the intertidal (Riedl, 1971; Riedl, Machan, 1972; r'1cLachlan,1979a) or the pumping effects of waves in the subtidal (Steel et al., 1970a; Riedl et al., 1972). In exposed situations filtered volumes may strip of average S-lD m3 per metre intertidal times on beaches. In the subtidal surf zone off open beaches wave pumping may filter and particulate organics in the sand. Under conditions, however, the high energy input of oxygen and the high drainage rates maintain and
oxygenated
. .
0
sand sand
representation in
et
of
a
0.1.,
( P.fter
1970; McLachlan
moisture content and surface temperature (Pollock, Hummon, 1971; r'lcLachlan et al. , 1977a) while in very sheltered situations it sharp can result in changes gradients, in chemical for example pH changes coupled to temperature IS78). a1. , vii th reduced layers three vertical zones (Fig. 2) may be distinguished, namely In sheltered beaches clean (oxygenated) sand, a gray (or RPD) transition zone and black (reduced) conditions (Fenchel, Riedl, 1970) . Horizontal zones may also be distinguished down the relatively constant beach from conditions of tbe the changes (Gnaiger et
this there
system are no
fully steep
vertical gradients in oxygen and oxidation states of sulphur or nitrogen et (Riedl, MacMahan, McLachlan 1969; al., 1979a). Permeability and flushing generally decrease with increasing shelter the until is drainage is so poor that sand constantly saturated.
vertical chemical gradients are then steep and reduced layer s occu r close to the surface (Fig. 2). The interstitial system cyclic changes related is to subject to tidal and
to the backshore 'VIithlarge and salinity fluctuations 1967; On greater Schmidt, exposed vertical 1970; beaches, extent
diurnal cycles and the seasons. In high energy beaches this mainly results in fluctuations in the water table, pore
325 the interstitial and better drainage, system lends itself to subdivision into strata. recognized
g ra ined (M d
Hummon in
t-1 ass
(1971 ) a coarse
tts
A TYPICAL
EAST
CAPE
BEACH
urn)
a c h use
beach which they could relate Sa Ivat's (1964) horizontal HcLachlan (1980b) strata on East Cape beaches urn) which closely parallel Pollock
g ]
Z 0 <
~
~
10 15 HORIZONTAL
---WATER TABLE
10 DISTANCE
20
(m)
30
40
50
60
and Hummon. In both cases the strata range from surface dry sand at the top of the beach down through moist
belo,,! tbe
layers to the permanently saturated zone water table (Fig. 3 and Table
of intertidal FIGURE 3. stratification based on interstit'ial sand moisture content during low tide for a beach in f-1assachusetts and the East Cape, South Africa. (After Pollock, Hummon, 1971; McLachlan, 1980). See also Table 1.
1).
Table 1.
Summary of stratification
water content (Modified from pollock, Hummon 1971; McLachlan, 1980). Moisture Content & Water
Zone!StratuFI
Degree of
Oxyger,ation 'Temperature
Zone of dry sandi :8ry sand stratum Zone of drying/ Dry sand stratum Zone of retention/ Moist sand stratum Zone of resurgence/ Water table stratum
High
Highly variable
High
Variable
Loses gravitational
water but
High
Little variation
r.1
d era
t e
to low
Permanently
saturated, little
Low to
very 10V!
Very stable
water circulation
326 Large numbers of microscopic organisms occupy the interstices: fungi, algae, protozoans metazoans. and Amongst the animals, metazoans that pass
bacteria,
cells per gram dry sand and increase with finer sediment and greater surface area (Dale, 1974; et al., t'ieyer-Reil
1978; ~1azure, Branch, 1979). Khiyama, Makemson (1973) recorded 362 bacterial strains in beach sand, many associated symbiotically wi th metazoans. Those on
through a lmm screen are meiofauna and the rest Individual dry weights are microfauna. 10-5_l0-8g 10-6 meiofauna, for 10-10 -IO-llg protozoans and for -1012g for bacteria (Fenchel, 1978; Fungi in sancJy Warwick, pers. comm. ) beaches studied are generally et concentrated a1., 1978). at A higher tide levels and have been little (Upadhyay flora of diatoms or flagellates only develops in sheltered beaches and is absent on very exposed beaches (Steel et This also al. , 197 Oa ; Brown, 1971a) . appears to be more characteristic of temperate than tropical beaches (Munro et ale , 1978) . In sheltered sands the diatom flora may be locally important and, due to mixing by wave action, may occur to some depth in the sand (Steele, Baird, 1968;
MeadoHs, recorded diatoms McIntrye,
1'1unro,
undamaged considered
sand grains made up 54-78% of the total numbers of 107_108 per gram of wet sand. Hicks (1974) demonstrated the presence of Azotobacter in Hla r in e sand beaches while Rheinheimer (1977) 31 strains from beach sand, of the majority showed salinity at 30-500/ gram and were 00 negative rods that cJecompose could proteins, carbohydrates and fats. which optima Andrews et al., (1976) studied microbial development in a model sandy beach, recording greatest abundance near the high and low tide marks. Organic have carbon and nitrogen been in the sand isolated
1977;
Brock
living to sand
bacteria grains
found to correlate with bacterial biomass and abundance (Dale, 1974; An d e r son eta 1 ., 1981; Bolter et 0.1., 1981) which can be up tc four times the 0.1. Liomass (1978) of (~1eyer-Reil, Faubel, procJuction a.t 15gc.m-2 the 1980). for a meiofauna Munro et microbial temperate
up
to
10cm below the sand surface and Round assemblage (1979) described a diatom living below the sand surface. Amspoker described the intertidal (1977) distribution of epipsammic diatoms on a
Californian permanently
estimated
beach. saturated
Hhere diatoms
sand is generally
beach and 72gc.m-2 for a tropical beach, the higher latter value due to greater water percolation. Mazure, Branch (1979) estimated an average bacterial biomass of weight in the sands Lagoon. disturbance
bacterial
Pearse et occur closer to the surface. 0.1., (1942) recorded Chrysophyta, Chlorophyta, Pyrrophyta and Cyanophyta on sandy beaches. Bacteria beach to the 1966, abundant and important in mostly being attached sediments, are sand 1968). grains (Meadows, range Anderson, 108_1010 Numbers
Dye
(19790.)
caused
greatly
activity beach sand. in et 0.1., (19790.) recorded numbers around 108 per gram down to more than 1m in the of an exposed beach. Dye
327 (1980a) was able to show fluctuations in oxygen uptake by tidal beach 1977; Small Fenchel, 1978; Hartwig, protozoans l1ave also 1982) . been
sand. Maximum uptake coincided with maximum water percolation through the sand and maximum fluctuation occurred at Desiccation of the higher tide levels. sand caused uptake or a severe bacterial drop in oxygen activity. Dye total
enumerated
on
ranged 101_103 per gram ury sand and they were found to more than 1m depth in a well oxygenated beach. Their contribution to total benthic oxygen consumption was significant, accounting for 15-25% of interstitial oxygen uptake.
r'1eiofauna
to partition
benthic oxygen uptake on exposed East Cape beaches and concluded that bacteria were responsible for most of the oxygen co~sumption except where there was a very t-1eye r -Re i 1 et rich macrofauna. microbial turna1. , (1980) estimated over time of 100h with daily microbial carbon production of 43I11g.m-2 in sheltered beaches in the Baltic. They concl uded that 50% of microphytobenthos primary production was fixed by microorganisms. Koop, Griffiths (1982) found a high bacterial biomass on an exposed beach receiving high kelp input on the
\,'lest
bave
received
considerable
(Delamare-Deboutteville, 1960; Swedmark, 1964; Fenchel, Riedl, 1970; Hieser, 1975; Giere, 1975; Fenchel,
1978).
Small beaches
meiofauna!
on forms
sandy
temporary
being Koop
bacteria
sediment.
that et a1. , (1982a, b) estimated bacteria were responsible for over 90% of these k e-l p s . of carbon utilization
of macrofauna, As or permanent meiofauna. we are mainly cJealing with interstitial meiofauna here, the temporary component is generally not of importance on open beaches. components The dominant sandy Leach meiofauna are nematodes of and
Annual turnover
(P/B)
of bacteria
in
this system was estimated (Koop, Griffiths, 1982; Griffiths, information the ecology 1983). we still of
harpacticoid taxa
individual
halacarid mites, tardigrades, gnathostomulids, hydrozoans and bryozoans. There is a well established relationship between the relative p roport ions of nematodes und l1arpaticoids and grain size. finer coarser mean Nematodes tend to dominate in sediments, harpacticoids in sediments and in sediments wi th size around et al., 300-350 urn they (Gray, 1971; Raffaelli,
of protozoans
la rge and qu ite abundant, espec ially on sheltered beaches with fine sands (Swedmark, 1964; Fenchel, Jansson, 1966; Panikkar, Rajan, 1970; Hartwig, parker,
grain
328 Mason, 1981; Hockin, 1982). Fricke, related reducing to differing tolerances to
1970).
generally
condi tions (Fenchel, Riedl, vertical distribution is shallower in the surf zone
is In sands above 200 urn the meiofauna usually entirely interstitial while
below 200 urn burrowing forms become increasingly important due to pore size restrictions (vlieser, 1959). Interstitial harpacticoids may occur down to t-1oore, 160 urn (McLachlan et a1., 1977b; can pursue an 1979a) while nematodes
interstitial
than on the beach but high numbers may st i11 occu r 10-30 crn into the sediment (McLachlan et a1., 1977b) . Dean (1981) looked at the effects of wave action on meiofauna abundance.
Horizontal cJistribution
may and
take
form
of
layers,
horizontal Riedl,
the zones,
125-100
Murison,
mode of life down to McIntyre, (Wieser, 1959; urn Fenchel, 1978) . 1973; In sediments nematodes As most open sizes in the nematodes always
(1973) optimum
Sjeograpbical 1970).
noncapillary
1972a,
b;
almost
l-1 r i son u
Itlhe r e 1979b) the 197 2a ; meiofauna is close to the surface and the distribution tends to take on a more
sands
of
230 urn to be
two-dimensional Seasonality
character
(Fig. 4).
for
faur;.a
the
of
suggested
meiofaunal
increases
of 250-350
bas Deen documented in several cases in temperate areas with the occurring lower meiofauna in abundance and moving deeper into the winter (Renaud-Debyser, in Harris, 1972b; Schmidt, 1972a, 1963; t.]i elk b,; Hulings, 1976; 1974; e, Faubel, Nodot, Feder, 1976; 1976; paul, areas 1980). warmer In seasonality is less clear (McLachlan,
sediment
On a vertical basis meiofauna distrito the degree of bution is related drainage and oxygenation of the sediment (see Fig. 3 and Table 1 ). Meiofauna abundance 1970; drops off drastically 1978) and in a the deep RPD and reduced layers (Fenchel, Riedl,
in sublittoral
cOIilplex
areas
(Emberton,
it may 1981).
McLachlan,
vertical distribution therefore only occur s on exposed beaches where reduced layers are absent (Renaud-Debyser, 1963; et
other than seasonal response Deen recorded as to have factors such as heavy rains (Bush, 1966), \-lavedisturbance (Boaden, 1968), tidal
r-1 i n eke, e
migrations
Different
factors
~Jest
(Rieger,
1 9 7 9)
ott,
and
1971;
he ide,
c h a n 9 e sin
moisture and oxygen over the tidal cycle (t-1cLachlan al., 1977a). et
defined
distribution
329
BEACH, INDIA
~
illillJ
All
5]
(b) A TYPICAL
INTERSTITIAL
FORMS
MANX BEACH
INTERSTITIAL) 3 EULITTORAl SPECIES (1 EPIBENTHIC 2 INTERSTITIAL)
MTl
harpacticoid copepods in a well drained, exposed beach (a) and a sheltered beach (b) where distribution patterns tend to be very three-dimensional and rather two(After (a) Munro et al., 1978 and (b) Moore, 1979b). dimensional respectively. all cases the copepods are concentrated above the low tide water table. On the In
sheltered beach interstitial fauna are mostly concentrated high on the shore where some drai~Qge occurs. Nevertheless zonation in this case is still mostly across the shore and not vertically in the sediment.
seasonality
are
becoming
clearer
as
non-selective
deposit
feeders,
result of ecophysiological studies on responses, preferences and tolerances of species (Jansson, important meioraunal 1967, Wieser, 1968a; 1975; Wieser Hartwig et al. , et al. , 1974; 1977;
feeders, carnivores and even feeders. studies have Various aggregation shown of meiofauna in response to specific bacterial strains (Gray, Johnson, 1970) or where there are high levels of organics generally or
localized
~lieser, Scheimer, 1977). A wide range of food items recorded for meiofauna
.
has
been
(Lee et
including detritus,
diatoms,
bacteria,
protozoa,
other
meiofauna
(Jansson, Fenchel,
Feeding
and dissolved organics 1969; 1968b; MCIntyre, 1982). 1978; Giere, 1975, categories bacterial include
Hogue, Miller, 1977b; Moore, 1979ai prey 1981") concentrations of or (Fenchel, Feeding 1978). rates have been estimated directly for a copepod (Rieper, 1978). have also lJeen 'l'he re growth, studies on ]?roduction and
330 population dynamics of important species (pawlak, 1969; Lasker et a1., 1970; Hall, Hessle r, 1971; Feller, 1980). Most round reproduction McLachlan 1977c; species have year times
i
adequate
considered
dissolved
and
ranging
1-3
months
(McIntyre, 1969
1975;
et a1., 1970). ~1easurements of benthic metabolism have attempted to partition oxygen uptake between these different of the interstitial fauna (Munro et 0.1., lS78; Dye, lS80a; 1981). Rates of mineralisation of organics lJy the interstitial system have also been measured in experimental sand columns in an attempt to understand these processes better and estimate nutrient cycling et 0.1., Boucher, 1970;
( r-1c In t y r e
Over the past decade ecological studies on sandy beach meiofauna have advanced from the more general surveys of the sixties. More attention has been paid to the complex of factors interstitial system and been bet ter (Gray, Rieger, composing the results have
components
analy sed stat istically 1971; Hulings, 1971; McLachlan, 1978; The basic pattern
Chamroux,
1976;
McLachlan
et 0.1., 1981c).
of meiofaunal response to different sediment types is well documented and to chemical features of the responses system are clearer. t-10re interstitial
ItlOr k
Vernberg, ratio
Coull
(1974)
estimated
the of as
is
needed
on
biotic
interactions
inc1udirg predation.
competition,
feeding
and
sandy for 0. sublittoral sediment and 1:0.54:0.04 tor an exposed Dye beach. (lS81) partitioned lJenthic
lJetween microfauna,
meiofauna
and macrofauna
rJith
regard largely
metabolism still
the
ratios
been
regarded
macrofauna.
partitioned Cape East
t-'JcLachlan
et
0.1.,
(19810.)
benthic lJeaches
interstitial
fauna
and
macrofauna
as
most other sediments, meiofaunal numbers fluctuate within an order of magnitude of 106m-2 and dry biomass values range 20-4400 mg.m-2. Some figures are compared in Table 2.
.
1:0.4. On a west Cape lJeach receiving a input, high kelp lJiomass ratios of macrofauna:
3.5:1:1.5 meiofauna:
bacteria
were
these 1:1:4.7.
ratios
the
because
these systems prov ides greatest habi tat availability and high percolation rates
two
uecades.
~Je have
yooc1
331
Table
2. Abundance and dry biomass values for meiofauna recorded from marine sands.
Source
Local i ty
Biomass
100 - 600 mg.m-2
'ilieser
(1960)
Buzzards Bay
Coull
(1970)
Bermuda
33 - 259 mg.m-2
McIntyre, Murison
(1973)
South Africa
McLachlan
(1977a, b)
South Africa
20 - 3 360 mg.m-~
')
South Africa
Fenchel
(1978)
sublittoral
50 - 1 700 mg.m-2
South Africa
Sten~on-00zey, (1983)
Griffiths
South Africa
:;eneral
idea
of
the
distribution
patterns
of
patterns relation
fauna in interstitial of features of interto many this However, in climate. of Riedl and wate r flow of
interstitial
meiofauna processes
and in
most
other
intertidal
This viater
through beach sand has not recei ved the The patterns water input, percolation and pulsing disturbance described by Riedl, Machan a considerable (Fig. 5 ) add (1972) degree of comprehension to distribution
flow is the superparameter controlling interstitial climate and the distrifauna in ueach of interstitial !.Jut ion uacteria and meiofauna sands. Both tend to concentrate in .the therefore r.1TL and below moist sand at and above the sand surface where there is bigh water minimum input, percolation and aeration, currents stagnation and pulsing
332
Under sheltered are not too severe. strata become the aerated conditions much flatter, the interstitial fauna concent rate s close to the sand su rface and the overall dimensions of the system take on a more two dimensional nature.
MAXIMUM
REACH
all organic it receives, materials whether these dissolved be or particulate. In this process bacteria most are mostly at food with
important,
the
top
of
chain. higher
There is little exchange trophic levels, especially beaches. of By the removal and materials
nutrients
OF STRONG
INPUT CURRENTS
mineralisation
of organic
inorganic
and the
micro- and meiofauna are responsible for the 'purification' of large volumes of sea water flushed through the interstitial 3. system.
-MTL
-MLWS
Hhile
as
quantitatively
MAXIMUM PULSING
-MHWS -MTL
-MLWS
and, in
l1igh
exceptionally
-MHWS
MAXIMUM PERCOLATION MINIMUM PULSING DISTURBANCE AND
The ir main feature on open beaches is the Ligh degree of mobility displayed by all species. These animals may vary from a few mm in length to lGcm and consequently a variety of methods have been used to sample them. Generally sand is passed through a O.5mm or lmm screen, screens though of 2mm in areas of coarse or even 4mm have sand, been
-MTL -MLWS
I MAXIMUM INPUT
MAXIMUM
MINIMUM
AERATION A ND
DISTURBANCE'
. .
~.
.. ...
. ... '.
-MHWS
..'
..~/. .
.
'
:,:<,:..
. '.
.' .
-MTL
used. Ci'hus not all .wor ke r s 11a ve sampled the same components of the community. The macrofauna community consists of
...::..<
-MLWS
FIGURE
5. Diagrammatic representation of
those organisms too large to move between the sand grains and generally in the size range lmg-2g dry tissue mass. Sucl1 species make up shifting populations in the intertidal and surf zones Hhen evaluating them of open beaches. it is best to consider the population across an intertidal strip of beach as a contracting and whole, this unit expanding as veach profile changes and
some important interstitial parameters related to water flow in an intertidal h~gh energy beach. (After Riedl, Machan, 1972). The interstitial system appears to be very efficient at mineralising virtually
333
tides
pass
Consequently
SHELTER
...
EXPOSURE .
;.:.: <;?.~.U
abundance and biomass of macrofauna populations consider metre wide will strips of beach and not square metres, stated otherwise. All authors unless should present their results in this way, at the same time stating the width of the beach. ~he macrofauna of sandy beaches includes most major invertebrate taxa although it has long been recognised that molluscs, crustaceans and polychaetes are the most important (Rees, 1939; ~iatkin, 1942;
South'\..;ard,
STAC EANS
~:tf:ti~t:i::!t:tt:;:::i:tti::;::::::::::~:::;~:;:::::;::::::;:;::::;:::;::;;::::;;;:::;;:;:;:;::;:;:::;;.:.;.:::;:;.;.:.;.;.
~
~
1952,
MOLLU
POLYCHAETES
Responses major FIGURE of the 6. invertebrate sroups to exposure the gradient on sandy beaches.
(Stephen,
1953;
SChuster-Diedricks,
Sourie, 1969,
1957;
Pichon,
1967;
is 1972) . a 'I'h ere / for crustaceans to be more abundant on tropical beaches or more exposed beaches and molluscs to De more exposed less and/or abundant on (McIntyre, temperate beaches 1968,
1928, Rapson, 1931, 1932; Holme, 1954; Figueras, 1954; Edgren, 1956; Ansell, 1961; 1959: Hade, MCIntyre, 1967; Smith, 1970; Brown, 1971b; 1971 ; Ansell et al., Irwin, 1973; de Villiers, 197 2b ; 1974; ~lcLachlan , Hanekom, 1979;
McLacblan, t1cLachlan
198 2b) .
Van et
cler
Horst, 1979b;
a1.,
1979: Lebe r,
and polychaetes are sometimes more these either of taxa abunoant than (Brown, 1981; 1983). t'icDermot t, Inoeed, Dexter;s (1983) work suggests the dominate most crustaceans that exposed beaches and polychaetes the most beaches with molluscs reaching abundance intermediate in ). (Fig. situatior.s In te nns oi 6 biomass, howeve r , molluscs are usually et important most ('I'revallion al. ,
ITiaX 1 ITiU;iL
Bro'vm (1982) re vi e'\-;ed i--lOrk on whelks or the genus Bullia and p~nsell (1983) has covered the genus Donax. There have been some general accounts of crustaceans em sandy beaches (McIntyre, Lopez, EpelcJe-Aguire, 1963; 1975:
Kamihira,
1979)
and
many
studies
sheltered
individual
species or groups.
have been extensively studied east coast of the United states 1971;
1967a, 1967b, 1968, 1970; Dexter, 1967, Sameoto, 1969a, 1969b; 1970; Craig, 1973; Croker
1970;
Dexter, et
McIntyre,
l'.nsell
1974, 1976; Eleftheriou, 1976; McLachlan, 1977a; al., 1972a; Bally, 1981; 1981).
Bousefield,
Shelton, Robertson,
et al., 1975; Scott, Croker, 1976; Holland, Polgar, 1976; Donn, 1980) and elsewhere (Williamson, 1951; Barnard, 1963; Fincham, 1981). 1971, IsOpods, have also 1974, been 1977; widely Kamihira, studied both air- and 1971, 1974,
~here have been many studies on the biology and ecology of dominarlt bivalves sandy beaches and gastropods of
water-breathers,
334 1979; Dexter, 1977a; K1apow, Fish, Fish, 1972; Kens1ey, 1972, Ferrara, Ho1dich, Brown, 1973; Johnson, Glynn et a1., 1975; Che1azzi, Ferrara, 1976b; Eleftheriou et a1. , 1980; 1972; 1974; ranges trom one (Gau1d, Buchanan, 1956; McLachlan et a1. , 1981b) to 82 (Vohra, this number decreasing exposure. divided beaches into exposure and, based 1971), with Bally (1981) three degrees of
increasing
Hendrickson, 1980). Mysids have been investigated by Fishe1son, Loya (1968), Brown, Talbot (1972) and Woo1drigde (1981) and cumaceans by pike, Le Sueu r (1958) and Corey (1970).
survey of 105 average numbers of species, abundances and c1ry biomass values for these (Table 3) . TABLE 3. Summary dry of mean macrofauna1 biomass values based
Bally, 1981).
Hippid studied.
crabs
have
been (1976)
abundance
and
Efford
on 105 beach
surveys. (After
distribution
in the genus
other studies include A1ikunhi (1955), (1969), Thomassin Matthews Di11ery, (1965, 1966, 1972), Efford Knapp (1970), Wenner (1977), Henner, (1979). (1979) and Subramoniam Fusaro Ocypodids have also been widely studied (Crane,
:' e 11 0 \.;' s,
High energy No. spp. Abundance m-2 Abundance m-1 Biomass g.m-2 Biomass 9.m -1 Sand particle diameter urn 11 400 20045 2.26 871 310
George, Jones,
Knott, 1972;
Hunter, 1978;
vannini, 1976; IS80c; Robertsor" Robertson, pheiffer, Pfeiffer, 1981; Hails, 1981; Hill, 1982; Wada, 1982;
~.'lcLa
chl an,
While with et
Ya z i z ,
1982
and
Ca in e
(1974 )
and
du
on
Preez There
(1981) have
have also
studied been
portunids.
studies
and abundance decrease (Angus, 1979; l'lcLachlan al. , 1981b) individual size increases, yielding high biomass values exposure
diversity
1982) ,
Laurence,
1982;
even at 10w"er abundances in some cases. Though most classifications of beach exposure are very subjective, the above trend s clear. highest remain The biomass values come from very exposed dissipative beaches, e.g. 6621 9 ashfree dry mass.m-1 in the Bast Cape (McLachlan, 1977a) and 25735g dry mass m.-l in Peru (penchaszadeh, 1971). These high biomass values are, without exception, due to filter feeders (Donax,
Dexter, 1975; 1977b) , a1., et (Clark 1962; Hock, 1981 ; Hilson, 1970; (New, 1981; Brown, 1982) and insects 1968; Craig, 1970; Orth et al., 1977; Griffiths, 1980; Chelazzi Stenton-Dozey,
Dexter,
Ebert,
polychaetes Longbottom,
et a1.,
1983).
Bally all
the
results to
of
surveys
1981.
Emer ita) . Exposed Leaches are thus not necessarily sparsely inhabited.
on a beach
l.1 c
1971 ;
l-1cLachlan
1977a) and may decrease again around the break point (McGwyyne, 1980) and then Day et increase offshore (Field, 1971; However, aL, 1971; Christie, 1976). on a beach dominated by kelp input, maximum macrofauna diversity occurred around the drift line at the top of the beach (Stenton-Dozey, Griffiths, 1983). ~he distribution of macrofauna along the beach is generally patchy, the comb i ned result of movement and sorting by the ash, or localized food concentrations S .,-1 species biological aggregations of (Loesch, 1976; 1978; Bally detail it to 1957;
AChutankutty,
Lawrence, Ferber, 1970; Bally, Albeit, 1981, 1978; et ale , 1981b; Lopez-Cotelo
et a1., 1982) beach slope (McLachlan et a1., 1981b) sand moisture (Hayes, 1977; Salvat Hithers, (Brown, 1964, 1966, 1967; Bally, 1981; 1977) 1964; food in the surf water Rapson, 1954; \jade,
1968; 1'1cLuskyet a1., 1975; Ansell et a1., 1972a; Nair, 1978; McLachlan et a1., 1981a; Hutchings et a1., 1983) and dynamic changes such as due to storms (Scott, 1960; Brown, 1971a). Effects of
d recJg i ng &nd beach nou r i shmen t have al so
Culter, ~redation
ale f
Hayes,
1977;
disturbance
Fusaro, 1980; Brown, 1952). (1981) investigated patchiness in on west Cape be a common beaches and showed occurrence ir! all
experimentally in a shel tered sand flat by Hoodin (1981). r--lany authors have monitored organic or chlorophyll levels in the surf without
correlations UncJ oubt 'v.ith high ec11:l
;
LJeing
with
0_ble to
faunal
hO\-lever,
species at all tide levels, often havins 101-102m. proportions of This sampling suggests transect that line
strategies
demonstrate
abundance.
1Jeaches
Liomass that on
must most
may if not
be
unreliable,
high it
inputs
of particulate
organics
particularly
replicated.
Dauer,
is
noticeable
Simon (1975), however, showed that by pooling the results of a whole line transect, error due to patchiness can De Hartnoll (1983) has red uced by 10-15%. quantified on a little size the species area relationship sandy beach. He found sheltered increase inc reased
tha t for
feeding very wbere rich filter populations occur, phytoplankton Glooms have !.Jeen recorded in the surf (New Zealand
Le'.;in et
Rapson,
al. , al.,
1954; 1981a;
Hashington
South
Sou
1979a;
.;:.frica
t h Am e r i c a
l'1cLachlan et
in diversity ubove
accu ra te
as
sample and
Gianuca, ~opulations
1983).
uevelop
Alternately,
0.2Sm 2
sampl ing
in
upwelling
sugge sted
an
of O. 5m2
is adequate.
resions, probably feeding phytoplankton acJvected shorewards upwelling cycles (HutChings et 1983) . Of the physical particle considered size the
Di
st
r i-bu t ion
on
the
beach
factors. macrofauna or
and abundance of organisms has been related to many Factors said to influence the include content sand grain size and/ 1970; (Longbottom,
factors, wave action and <jenerally been have most important although
organic
336 Eleftheriou, Nicholson (1975) show that grain size alone can not characterize a beach. As particle size, beach slope and wave action are closely related, defining the former two fixes the latter 19800.) . (Davies, McLachlan, 1972; et McLachlan significant macrofauna
( 1981 b) ale , correlations
20 (a)
4000
115 en LU C3 LU g, 10 u. 0
II) 5
3000
I I I I" E
2000
obtained between
LU
a:
e"
~eC1'
~~~/ /
LU II) :J 1000 Z
:E
en a:
l>'~ flJ/
"e~
and both grain size and beach slope, but not wi th wave acti on est imates for beaches on the south and east coast of South Africa (Fig. 7). This suggests that it is not wave action but rather steep slope and coa rse sands that 1imi t the fauna. very exposed beaches (in terms of wave heiCjht) often have richer faunas than less exposed beaches where the latter have coarse grains and steep slopes. Finer sands result in flatter slopes as does heavier wave action. 'i'he flatter the slope of a beach the more
eve r: y 1 wa v e e n erg y
:J z 0
C11 ~(;>o/
c:,Q
-""
(\J" ~.Y~ 0
4'0
~o
BEACH
~o
SLOPE
\0
4000
20
(b)
115 en ~LU C3 LU g, 10 u. 0 LU co 5
~:J a:
\' 3000 I
I
E
2000 en a: :E
LU co
:J 1000 Z
is dissipated
in the
surf zone and intertidal and this is the Thus a very exposed crucial factor. beach can support a rich mac rof auna if opposed as to dissipative it is relective McLachlan, (e.g. 1977a; Maitland
Copali s beach,
FIGURE
7.
TIle species
relationships diversity
be t.,leen
macrofaunal
anu
abun-
beach,
Lewi n
dance
Indeed, high energy et al., 1979a). dissipative conditions seem optimal for the development of a high lJiomass of
and the slope (a) and particle size (b) of ueaches in South Africa. (After ~icLachlan et al., 1981b).
filter feeders. Much less is known of macrofauna ecology intertidal. Fauna is below the generally absent around the break point but increases in abundance
onshor e and
1971, Dexter, 1978; 1970, 1972; Hill, Shin, Hunter, 1979; 1981, 1982; Kastendiek, 1982). The zone inside the break point bas been termed the inner turbulent zone and the area outside this (out to where wave effects on the bottom
and diversity
wi th
are
zone. Masse
negligible)
the
outer
turbulent
offshore
in
accordance
less
turbulent
10031 m-2 and biomass 2.5-11.7g 1048 dry mass m-2 at various localities in at depths 1,5 Sm. the ['ledi te r ranean
McIntyre, Eleftheriou
(1968) recorded an
337
aver-age
dry
intertidal
subtidal
biomass of 1. 3g .m-2 in the the and 3.7g.m-2 in with 62 species on the beach The
the midshore in New Zealand. In South scheme, Africa with Dahl's rnodifications, has been found acceptable
(~lC Lach lan,
and 116 off the Scottish shore. poorest zone was just below LHS.
t-1uch has
1980b;
McLachlan
et
0.1.,
1981b)
been
written
on The was
serious attempt
at providing
I MIOLITTORAL
II
RETENTION
SUBLITTORAL
FRINGE
of (1952) (Fig.8). very Dahl This was suggestion by to an earlier similar Davenport (1903) . Based on the distr icrustaceans bution in northern of temperate and Sou t h Am e r i can be a c h e s he proposed essentially three zones equivalent to the three zones of rocky shores. These zones were: fringe characterized subterrestrial ocypodid warm areas crabs in amphipods in cold areas; tal it rid mid1ittoral zone characterized the by cmd the by
. .
.
B~<6~~~S
~~~~~
ELWS
FIGURE
8. Diagrammatic
representation zona t i on
on
of
cirolanid isopods (although these may be absent in cold temperate areas); and the fringe with a mixed fauna Sl.:blitt.oral often characterized Ly l1ippid craus in
the tropics and haustorid
sandy
1967) Dased
and
other
alternate
scheme
on
described an physical
amphipods
factors. He clelineated four zones on the ueach, Lased on water saturation B) . (Fiy. Pollock, Humon (1971 ) used were studying, exce~t that they suudivided the zone of dryinSj and they extended L,ones strata their into covering the 'whole interstitial system 3). Salvat's Table 1 and Fig. system bas Deen found reliable in describing zonation of macrofauna, (see particularly isopods (Hithers, Bally, 1981). Both these 1977; authors this they system for a Massachusetts beach
authors have tried varying results. scheme with tbis Gauld, 3uchanan (1956) round an overlap on west of talitrids and ocypodids Subsequently African showed (1972) Singapore found
et 0.1.,
Excir-olana
recorded shores and also Lower zones in this zone. correspondence. Philip less India, Vohra (1972) in and Jaramillo (1978) in Chile
in Dahl's
scheme useful and Escofet. found it fairly suitable Uruguay and beaches in Brazil, for J.~rgentiria . Hood (1968), however, found circolanids replaced by a sphaeromid on
(1978)
admi tted, lloweve r, tho. Doundar ies t are not sharp and that oones Sjrade into each (1976) otber. Eleftheriou, Jones pointed out that differences in inter-
338 tidal !..Jetween distrib'.ltion different areas and species make eurydicid isopods unreliable indicators of zonation as proposed by Dahl. They did not, however, attempt to use Salvat's scheme. Views on zonation of macrofauna on sandy the larger one in the eulittoral (e.g. Amphidesma spp. in New Zealand, Rapson, 1952; Donax spp. in India, Ansell et a1., 1972b; Donax spp, in the Last Cape, MCLachlan, 198Gb). Besides level, patterns at intraspecific zonation
i:.onation
community
beaches are thus highly conflicting, a situation not surprising in a dynamic environment wi th a highly mobile fauna. It may safely be said that zonation, in sense, has never the classical been proved on a sandy beach, i.e. sharp boundaries have not Further, individual clearer zonation than whole. Indeed, Brown of the opinion distinguishable breathers breathers been demonstrated. species show much the fauna as a is are air (pers. comm.)
has Leen in several species. his '1' takes the form of L-.onation of and bas ueen recorded in typical of open beaches, crustaceans al. ,
\J 001
classes
1975;
g e ,
d rid
1981; 1982;
Ansell,
1980;
Brown,
Haley, lS82).
in the supralittoral and water below them. However, two boundaries are clearly and indisputably open !..Jeaches, namely on the visi!:Jle drift line and the edge of the saturated sand where the water table reaches the to Dahl's surface. These correspond although boundaries. Howeve r , the boundary between Salvat's zones of retention and resurgence are not obvious considerable evidence that is there meaningful midlittoral These beaches. on zonation schemes are Within these summarized in Fig. 8. zones most species sbow subzones and subdivision made can be of the many
differential sorting of the sizes in the swash or to active miS:]ration to areas differentially suitable to the orsanism at different life stages. Zonation as the researcher records is, however, only the uistribution it
of the t!sammolittoral fauna c.L an instant in time and most species undergo tided migrations of some sort. Ty 1?i call y t his involves a simple movement up and dOl-ln allowins the animal to stay in the swash zone Y/here conditions for feedins are opitmal and predation minimal. In molluscs this movement involves rio endogenous rhythms but simply a series of responses to changing lJhysical conditions, most
the tides
the
th !..Jeach ,Ii
there is often blurring of boundaries. Isopod species usually exhibit a clear zonation near the top of the shore and midlittoral (Withers, 1977; Eleftheriou, Jones, 1976; Bally, 1981; McLachlan et al., 198+b; Dexter, 1983). Bivalves, often with two also exhibit zonation, species occupy ing a beach, the sma ller species in the sublittoral fringe and
notably
thixotropy Turner,
the
degree
In some molluscs' it may be TIIOre complex, e.g. Donax serra in the Last Cape, where
Trevallion,
1971).
there is not normal tidal migration but rather semilunar movement up and down the shore to occupy a position near mean
339
tide
tide 1979c)
level during
and to been
low 0.1, of
compress
the may with be
so
that
at
high into a
tide narrow
most
of
U1cLachlan
invertebrate com~ressed
population
on the beach
strip between to into
overlap
.
these and to night as behaviour 1963, 1965, shown at springs rhythms more include (e.g. are complex. entry mysids, increased opposed (Papi, 1972; to
species.
tidal
audition
normal
the
movement
endogenous have been plankton and during and 1963; orientation Enright,
subtidal bas been recorded in winter Donax parvula (Leder, 1982b). Temporal changes in Deach
in
the
macrofauna
anphipods
isopods),
uramatic
are those
assoc-
0.1., 1968, 1969; Cubit, 1969; Fincham, 1970, 1973; Alheit, Naylor, 1977; McLachlan 1976: r'1acQuart-110ulin, et 0.1. , 1979c; Harsh, Branch, 1979: Hager, Croker,
et 1981) activity it as prevents the tides
Hammer
et
iated with the monsoon in India \lhe r e most of the fauna Gisappears during this period (Ansell et 0.1., 19720.; D\JiveCli et 0.1., Longer years 1954;
1978)
et
0.1.,
1975). several
1980;
0.1.
Fusaro, 1980;
advantage
Jaramillo Ugolini,
increased that s~randed
1980; Scapini, of
is being
term fluctuation
spanning
springs
.
may
Dramatic
animals
Ansell,
Trueman costs
(1973)
calculated in
the
form of occasional mass Ttlortalities, e.g. Donax spp. (Orton, 1929; Fitch, 1950; Loesch, 1957: JOhnston, 1966, Nel, 1968; Grindley, 1968; de poi to most
sorJous
term
also
take
energy
it ,,;as more Emerita and ?~ofitable for the animals to migrate than to attempt to maintain position. behaviour has several Tidal migratory
Dor:ax and
Villiers, 1974) such as related uinoflagellate blooms. been and reported in for several
communities.
advantages: it allows animals to stay in the zone optimal for feeding, i.e. the edge of the swash zone; it keeps
thern
. ~J.
an
unstructured
and
physically
too shallow for many fish predators but just immersed enough not to be fully open to birds: it helps to prevent animals being supralittoral enables them stranded
for m s ,
like a sandy beach, to be unspecialized (Brown, generalists with board niches Talbot, 1982; McGwynne, 1972: Brown, 1980;
the Bally, 1981). Because of this, physical
on the shore; in
e .g . Ty 1 0 s , it
nature
mobility amongst unlikely, of LJe
of
the
the
environment the
sandy
to
move
d9wn
the
beach
of
fauna,
beach the
though competition
migration
on
naturally
and
has all
marked
exploitation
ef fect s
zonation
zones
ruled
out.
Hilson
(1981)
340 demonstrated negative interactions in a dense assemblage of deposi t feeding polychaetes, but this was in a very sheltered beach. Trophic include feeders groups among the macrofauna predators, filter and, on less and
.
predators.
As the bulk of the fauna are usually filter feeders, the size of beach populations is probably closely related to the richness of inshore waters in particulate organic material (Ansell et al., 1972a; Hutchings et al., 1983). In some areas large amounts of mac rodebr is may cast ashore, particularly ue adjacent to kelp beds. In such situations vast populations of supralittoral scavengers develop (Hayes, 1974;
Griffiths, Stenton-Dozey,
exposed
shores, feeders
feeders. dominate
Filter
consist Donax
mostly
bivalves
The
tellinaceans 0f
of the
high
1983).
in sandy
Deposi t feeding
common
1981).
except in very situations. re sheltered In cases \vhe the fauna is impoverished, however, such steep very coarse, beaches, as on supralittoral scavengers may dominate as a result of the virtual absence of other species (Gauld, Buchanan, 1956; Dye et al., 1981; Hoo1dridge et al., 1981). On
shel te red Arenicola, beaches deposi t and feede rs, e. g. Callianassa Scolelepis,
In other areas carrion input may be high, e.g. in the form of cnidarians, and in such cases midlittoral such as Bullia reach high (McGwynne, 1980; Brown, 1982). food webs scavengers abundance
have been given for beaches in South America (Koepcke, Koepcke, 1952; Gianuca, 1983), the Cape (Brown, 1064; Griffiths Peninsula et
t'Iacrofaunal
may be very important or even dominate (e.g. Saloman, Naughton, 1978: McDermott, 1983) . The tropic structure of the beach macrofauna is therefore normally dominated by mobile filter feeders. be more forms may feeders usually Even where other abundant, filter dominate biomass McIntyre,
The main predators on the macrofauna as listed by several authors are birds, fishes and crabs (Coe, 1955; Loesch, 1957: Masse, 1963:
.
the
E;ast
Cape
(McLaCfll
an
Hade, 1975).
1967;
Penshaszadeh,
Olivier,
Eleftheriou, (Dexter, 1979: 1976). P.s there is little or no primary production on the beach, the macrofauna is dependent on food imports ad jacent systems,
The former two v/ill ve discussed being later. Crabs, the tJart of benthos, represent a form of predation resident in the veach/surf system. Du Preez (1981) has made a detailed study of the swimming crab Ovalipes on East Cape beaches and shown it to lJe an important preda tor on Donax and Bull ia. Seve ral authors Iilentionthe importance of both ghost crabs and swimming crabs as predators on ueaches (Loesch, 1957; Koepcke,
Virnstein 1953; (1977)
from
sea (Brown, 1964: Ansell et al., 1972a; McLachlan et al., 1981a: Griffiths et the most is sea The al. , 1983). important filter and and and supplies for particulates and scavengers for and deposit feeders carrion
other
Ansell
showed
et
al.,
1972a).
the
importance
macrodebris
of
crabs
in
controlling
macrofauna
341 communities sand in gastropods burrowing by predation in subtidal Chesapeake Bay. Naticid are important predators on but are not generally 4. PhYTOPLANKTON Blooms of diatoms persistent a typical or sporadic, feature
in
now
surf sandy
zones, ueach
appea r to ue
bivalves
uf many
found on open ocean beaches. mortality on sandy This includes not only mass poisoning (de due mortalities to being cast Villiers, 1974) but also
"natural"
Besides
predation, times
surf zones. Rich surf phytoplankton blooms have been reported from the Hashington coast, U.S.A., (Thayer, 1935; Lewin, Norris,
,
can at beaches.
be
important
1970),
above the shore by storms (Brown, 1971a; Penshaszadeh, Olivier, 1975; t:1cLachlan Young (1982) et a1., 1981a). McLachlan, suggested the importance of upwelling temperature drops sudden in mobility as a factor contriretarding to..; a r d s this. Ansell et a1., bt.:.':ing and (1978) mortality that have shown where rate 1S fairly constant,
coefficient,
the (Gunter, U. S. A. 1979; Gunter, Lyles, 1979), New Zealand (Rapson, 1954; Cassie, Cassie, Lewin, 19GO; Norris, 1970), South Africa (McLachlan, Lew in, 1981) America and the east coast of South
(Gianuca, 1983). These blooms are mostly cor.lposed uf masses of cells LJelonging to the of one or tvJO species
genera
Chaetoceros,
Asterionella,
Aulacodiscus The
anc1
the blooms
Gunter,
are of
related.
cornrnercial
dominatecJ
strips
hugging
the
ueaches.
cases
by
Eeavy
rains
leach
nutrients
from
the
or Donax
..7h ere
heavy
mortality
may be
Ilian.
land and these, lJeing retained near the beach by calm seas, apparently cause the blooms. These blooms take the form of strips S-6m \JicJ for miles along the e shore. All the other reported surf zone blooms occur as patches uf phytoplankton
~wi thin
distribu~ion and diversity of beach macrofauna by physical largely determined is prime among which factors, are wave
l~
rnay~
be
concluded
that
the
~well
c:levelopec1
surf
zones.
action,
particle
size
and
beach
slope.
these, have
those
uff
the
~Jas hi ng ton
Of coast
Food input probably determines abundance or both scavengers and filter feeders. most pronounced feature The clegree macrofauna animals is a bigh mobility and tidal migrations beaches. macrofauna The on exposed of main role the is to process food imports,
been
most
intensively
studies
342 retained
often clearly retention
in
the
surf on
to the long tion
more
has
been
to the ecology
!Jlooms. described zone in in Jijina, physical relation
distribution
(in prep. of at
of
a, the 13
features
blooms
(1974)
species Aulacodiscus
coastal beaches
and
on
Oregon.
long flat
littorii
dominant
Blooms were
beaches beaches
armatum dominant 1950 and Chaetoceros after 1950. Other studies looked at light utilization (Lewin, Mackas, 1972) , nitrate reductase activity (Collos, Lewin, 1974), daily periodicity (Lewin, Rao, 1975), C:N ratio (ColIos, Lewin, 1976), respiration (Robertson, Lewin, 1976),
a1
.,
while steeper beaches and near rocks has poorer blooms. were Low air and Hater temperatures associated increased diatom with densities were
densities.
while
inversely They
chemical
and
composition
the cIa y co at
(Lewin
( Le win
et
et
of
beach
and
characterisin bloom
1 9 7 9 a)
tics
were
diatoms
development.
f.1cLachlan
suggested
trap lJeach out that
Lewin et al., (1975) analysed envi ronmental conditions associated with the finding two years, the blooms over populations relatively constant diatom (1978) Lewin over this period. evaluated surf. seasonal Nitrate concentrations fluctuated widely and in relation to season, upv/elling phytoplankton growth and ammonium seemed to be an important nitrogen source, during summer. particularly Lew in et controlling factors cycle of nitrate in the the
form
circulation
develop
nutrients,
and on
pointing
this
is
most
likely (i.e.
long
beaches
vf
flat
slope
1.1cLachlan!
I-.ewin
of Anaulus birostratus from the Bast Cape, South Africa, and shov/ed tha t the blooms were positioned over rip currents by <.lay. All surf bloom torming species
show a daytime buoyancy and this results
ammonia
that
excretion
the razor
in tbe collection of diatom cells as a foam or scum over rip currents. Presuffiably the opposing forces of incoming waves and outgoing rip current keep them in position. ~Jhile c.:iiatom cells may travel all around the surf cells, the rip current area bottle-neck
f.1cLachlan, blooms in
populations
regenerate
of
Washington
beaches
of Le win
significant
amounts
zone. into the surf ammonium (1977) summar ised knowledge of blooms, pointing of the Diatoms
circulation
may act as a
these
for
Lewin
buoyant
(1981) in
material.
also not nutrient showed that
diatoms and
are the main
clams while the clams in turn regenerate inorganicnitrogen which is utilized by the diatoms.
developed is low,
water
very high
concen-
nutrients.
Though
tration
inorganic
supply
is are
constnat
as
nutrients
continually
343 draining out of the beach. Nutrients species commonly occur in well developed
are generated on the beach both by the filtration of water and mineralisation fauna of organics by the interstitial (McLachlan, 1980a, 1982; McLachlan et al., 1981a) and by excretion of the macrofauna
(Proscb,
~1cLachlan ,
surf zones on gently shelving beaches. Diurnal buoyancy of the diatoms results in their concentration over rip currents by day. Nutrients are derived from many sources, l..>u the beach is probably most t important. Inorganic nitrogen generated as ammonium
l..>y
1983).
and Cape beef!
Both of these
sources
are important
macrofauna
and
have been shown to generate East amounts of nitrogen on beaches. Subsequently wind has
significant
shown to be of overriding importance in controlling sbort term changes in blooms in the surf (Romer, Sloff, pers. comm.). 2stimates of primary production in surf
as nitrate Ly the interstitial fauna is constantly draining the into surf and passing through rip currents where the lJlooms are situated. In this way the lJlooms are positioned in a site of steady nutrient influence controlling circulation
more of the
zooplankton
excretion
and
supply. on
.wave
Hind Lloom
zones off beaches have been made by (1960) and Edwards CaSSle, Cassie (1973a). The latter obtained figures of 10495 and phytoplankton off shallow beaches. productivity blooms
Schaefer,
action
patterns. interaction
and to
surf know
wind,
between
waves,
Estimates
and blooms. \Je also need more reports on the sources of primary production in surf zones that do
f!utrients
are
figures
a1., (1982), Hilce, and Quinlan et ale (1983) Quinlan (1983) free living reported masses of on algae, Pilayella, sand on oall-liKe
ReceDtly h1ilce et bottoms and in the surf off beaches in This species Nahan t bay, Hassachusetts. peak and biomass in summer reached by fragmentation and reproliferated generation. uncertain
.the
from the literature. papers Hhile several taxonomic list species taken off sandy beaches, few were sampled in the surf zone or include ecological notes. Brattegard (1969 and subsequent \,orks) described mysic1s from shallow vlater off the Bahamas including some from sandy habitats and similar accounts are given vy Higley, Burns (1971) for the Atlantic coast of the U.S.A. and Bacescu (1975) for Tanzania. Bowman (1971) described coast a by and of copepods the uff the and one lay
zone
zooplankton
are
but
beach/surf
system
is
considered
important.
~... .J.L.
'coastal
one'coastal'
association'
may
be
concluded
diatom
blooms
'estuarine'
species.
Outside
'shelf'
'oceanic'
associations
344 by dominated respectively. studied four and Sander, and seven Moore species (1978) copepods outer limit of distribution near the outer limit of rips. He postulatecJ that detrital foocJ in suspension c1ecreases outsicJe tl1is zone and that zonation was mainly related to nearshore, tl1e wave-induced Several circulation ~atterns. have recently been clone
inshore
offshore
around the West Indies as shallow as 10m and recorded average numbers of 100500.m-3 with other groups making up 11-22% of Youngbluth total zooplankton numbers. (1979) studied zooplankton
studies
within lkm of the coast of puerto Rico at about 10m depth. He recorded numbers of 41-7568.m-3 of which 65-84% were copepods. Settled volume biomass values averaged 0.086ml.m-3 and highest numbers were recorded at night. Moran (1972) found the benthic mysid, Gast rosaccus sanc tus, to occu r in shoals along the shore in the top lcm of sand, but moving into the top I-2m of water at night
exhibited
on surf ~one ~ooplankton in the Last Cape. In IIIOst of tl1iswork the use of bas attempted to avoid the such as normal loss of large forms mysids, a problem often not appreciated by workers usins small nets. Cockcroft nets (1979) sampled inside the breakers at 1m depth, just outside at 3m and further out at beach. outside
efficiency Abundance
large
breakers
netting .surf).
in
the
Mediterranean.
It
\-las very
(1977), and an
ancJ
after dark when dry biomass values ranged 6-980mg.m-3. Mysids dominated biomass
increased
two in
benthic
the
mysids
ran.ean,
i.jedi ter
mean
mysid
in
benthic
(21% at 1m, 83% at 3m and 8496 at 10m; 63% ) hith copepods averagins only At site prawn, 13% . the the 1m
africana,
\Jas
patterns.
similar planktonic
Macropetasma
very The
impor tan t althoug h poo rly dominant species overall Mesopodopsis slabberi.
distinctly l1igher tban
sampled.
crustaceans, activity of benthic mysids, amphipods ancJ particularly isopods, is .well known (Alheit, Naylor, 197G) . The most detailed account of su rf zone zooplankton yet published is the work of Clutter (1967) at La Jolla. He studied zonation of four benthic the nearshore and five pelagic mysids out to 17m depth. All species clear zones, bottom. A
species,. where rip
offshore
Bay
and
about
the
same
as
local
estuaries,
indicating
richness of zooplankton in the surf ~one. Subsequently detailed occurs zones. this Cockcroft (1983) has macJe a which
stucJy of
Macropetasma,
formed
schools
and occupied
prawn is really a nektonic form. exhibits clear zonation with It a juveniles inshore ancJ adul ts out as far as 20m ciepth. by day They and shoal bottom disperse near the at night,
345 being most abundant inside the breakers. Numbers recorded averaged 21-37.m-3 giving a dry biomass of 0.9-1.8g.m-3. This species uses the surf zone as a nursery area and, being an opportunistic omnivore, it feeds on detritus, phytoplankton and small crustaceans. Shoals tend to move inshore at night and to concentrate around phytoplankton
blooms by day.
and even
reached densities exceeding 15000.m-3 (mainly juveniles, dry biomass I.02g.m-3) behind the lJreakers at mysid species seldom Other 10.m-3. Dry more than zooplankton biomass averaged about 0.25g.m-3 behind the breakers and 0.015g.m-3 offshore, an order of magnitude lower. times. totalled Mesopodopsis virtually moving abundant population juveniles even into formed vast shoals which
(pers. Eomer comm. ) is food chains around surf blooms, including small
zooplankton.
He finds that zooplankton is concentrated around blooms by day but more dispersed at night, this resulting in a slight decrease in biomass after (Macropetasma, Crustaceans dark. mysids, of
copepods)
to deeper
water)
though
for about
85%
estuaries.
biomass
siphonophores,
others for the total numbers :emainder. in a bloom during the day are about 2000 zooplankters.m-3 with a dry lJiomass of "') higher values, up However, O. 8g .m-.).
cbaetognaths,
~1 e a n
a mysid dominated zooplankton cOIllTIlunity of very high biomass. Abundance and biomass in this region are an order of magnitude higher than offshore and even higher than recorded in most estuaries. Hany have probably underestimated the contribution of the larger and more mobile zooplankton through using small nets. Clearly our knowledge zooplankton is abysmal. beyond species lists
abundance. extrapolate
\Jb i 1 e
_':/
to
750g.m"", swarms
have
been
recorded
dense
of plankters,
in particularly
authors
where prawns were abundant. Hooldridge plankton of (1983) Algoa studied Bay from the zoojust behind
the breakers to about 4km offshore (20m depth) using a net of 1.5m diameter. y;h salps, medusae, ctenophores and i I e others biomass. accounted crustacean were often making Fifteen common, up species crustaceans 80% of of mysids dominated, about
it the
from
hard uata currently available, three points seem clear: i) the zooplankton of
exposed su r f zones may be ve ry r i cll wi th
high dominate
are
lJiomass
values,
ii)
large"
in importance. A single species of mysid, Mesopodopsis slabberi, was by far the dominant with species, average especially numbers close around inshore,
crustaceans
mysids, penaeids) the biomass (as they can cope of these areas) but are to sample, iii) the re
(e.g.
346 clear offshore zonation patterns and diu rnal changes in abundance. Huch wor k remains to be done in this field, in particular relating distribution and abundance to wave induced circulation patterns and available food in the nearshore zone. 6. FISHES studied better than Though far zooplankton, we still know comparatively little of the ecology of fishes from surf zones off open beaches, largely because of the difficul ties of sampling pearse et al., in these environments. (1942) list fish caught by seine net off beaches more Carolina and North extensive Gunter
( 1 963 ),
feeding subtidal
of off 1966;
Corlett,
Edwards, steele, 1967; 1968; poxton et al., 1982) . Lockwood, 1974; Among these fish, plaice (pleuronectes platessa), dabs tLimandia limandia) and
turbot (Scophthalmus maximus) are the most important. summer, after In large numbers move into metamorphosis,
fin e san d bay s
'vI
her e
the
wa t e r
is
1e s s
than 3-5m (Jeep. There they feed on demersal plankton an<.1 benthos, including mysids and the siphons of Tellina and tentacles of }?olychaetes. Horta1i ty is high and the populations of these juveniles through deeper winter. United (1958) In the States Gunter sampled surf fish off the 'l'exas coast F recording fewer species off open beaches than inside tbe D3.rrier. He mostly caught small and juvenile fish which spring were most and aLundant in summer. McFarland (1963) zooplankton 'v,asthe primary for the Texas. found that food source generally autumn water until again at decline they the rapidly move onset into of
reports et
of al.
( 1967
fishes
in
surf
are given by
1970 ), Cup k a
to juvenile has been paid as nursery using beaches flatfish grounds (r1ace r, 1967; Edwards, Steele, et a1., 1970; Steele, 1968; Edwards Edwards, 1970; Steele et al., 1970b; Jones, 1973; Gibson, 1973; Thijssen et al., 1974; Lockwood, 1974; Pox ton et al., surf the 1982). In South Africa Rossouw (1983) has studied elasmobranchs from zones with particular emphasis on
abundan t trophi c 9roup and even mullet t (Mugil cephalus) and benthic feeders took large amounts of zooplankton at times. He recorded 47 species with an average (;f four in winter (biomass 25.81b/acre) an(J 16 in summer (biomass
annulatus, sandshark Rhinobatos Lasiak (1982, 1983) has made a deta ile(J
investigation
of surf
zone ichthyofauna
communities and Romer, (pers. comm.) has investigated food chains around surf phytoplankton blooms. In the cold temperate and Europe shallow
impo r tant as nu r se ry
103.21b/acre).
High
primary }?roduction
in the surf, he concluded, was more than suffici"ent to support the fish community. MOddle, Ross (1981) also recorded midwater planktivorous fish dominating the surf zone in the northern Gulf of Mexico.
Armitage,
and
several
authors
have
studied
Alevizon
(1980)
347 recorded the diet of the Florida pompano (.Trachinotus carolinus) and found Donax to be an important item. et ale , (1960) recorded 71 Carlisle species off Californian beaches with an average of 284 specimens per beach seine surfperch, Amphistichtus The haul.
argenteus,
Edwards (1973b) sampled with a beach seine net off two Venezuelan beaches and recorded 49 species (31 at one beach and 32 at the other) which he di vided into plankton feeders, fish carnivores, benthos feeders and berbivores estimated unpolluted a at (Mugil). 5g.m-2 and beach. beach Fish biomass dry mass at 19.m-2 at was the at
and
the orbina,
Henticirrhus
polluted
undulatus, were dominant but flatfish, skates, rays and sandsharks (Rhinobatos) were also common. Best catches were They made a taken on the low tide. detailed study of the biology of the surfperch including reproduction, growth and feeding and Emerita analoga,
the diet.
Plankton feeders and benthos feeders most were important. Penchaszadeh (1983) found mysids and other motile to be the main prey of crustaceans fishes off Venezuelan beaches. In Australia described the Lenanton utilization et of al. surf (1982) zone
accumulations lif macrophytes and their associated amphipods as nursery feeding the the areas for species. In the 0+ year classes of three fish
and mysicls were the dominant foods though fishes, particularly anchovies, were During sampling important. also 71 species of fish were recorded.
(1983)
Schaefer (1970 ) studied feeding of striped bass, Morone saxatilis, in Amphipods Island. surf at Long
Africa, Rossou\-J has studied elasmobranchs with particular reference to the biology
(1983)
Eastern
Cape,
South
of
the
sandshark,
Rhinobatos
annulatus.
f-';cDermott
sampled
the fauna of a of
b irth
This species comes into the surf to give ins u mm e r. Juveniles stay in the surf but adults
on
barrier beach
r,love
Iilysius,
offshore
small
in
Donax
cores and seine netting. The dominant benthic organism and main food for the polychaet, Scolelepis the fish was squamata, which reached densi ties up to in a 40000.m-2 (=50g dry mass.m-2) band approximately 20m wide. Of 26 fish species only seven were common and one This species, Menidia menidia, the silverside, made up 85% of fish of numbers and 40% fish mass and of its diet. Scolelepis formed 70"-80% (1982a) recorded four fish species feeding on beach macrofauna, including
Leber
winter.
and
'.I.'heyfeed
various rays, other elasmobranchs, duckbill Hyliobatis spp., come into very shallow water to prey on Donax spp. The se ,
lither benthic prey.
Amongst
together
wi th lither large
As
resident.
in the case of flatfishes in I..;uropean waters, the <jreat success of skates and rays in these environments is probably due them to to their forage flattened in very shape shallow allowing water,
the silverside.
thus being able to take even the macrofauna that undergo tidal migrations.
348
investigating phytoplankton
controlled by winds. six feeding categories, narL1e 11' : benthic feeders, planktivores, uetritivores, }?iscivores, herbivores and omnivores.
Herbivores
blooms. He has found that mullet t, Li za richardsoni, grazed directly on the phytoplankton and are taken in turn by large r predatory fishes. The mu llet t as well as other juvenile fish in the surf also prey extensively on zooplankton. Lasiak (1982, 1983) has made the most study to date of surf fish detailed seining on two Algoa Bay communities, beaches At over a period of three she years. recorded the less exposed beach
Lasiak
(1982) distinguished
to the presence of
some rocky reefs r,ear the sampling site at the less exposed beach, and were virtually absent at the more exposed beach. The mullet, Liza richardsoni, was the only detritivore, but in Lasiak's study fed mainly on zooplankton and may thus be considered a planktivore.
}ler simplified
66 species and 29 at the more exposed numbers site with teleosts dominating dominating biomass. elasmobranchs and Seven common species were resident while species intermediate were or 0ther sporadic in occurrence. The community showed no overall seasonal trends but a degree very high of short term variability which seemed to be la.rgely coupled to the effects of wind on the biological zone as several surf to parameters were closely correlated wind and intensity sampling. Studies over 24h clear trends though maximum
direction
planktivores,
may
At both 0eaches Iilotile organisms (mysids, prawns, fish, cephalopods and zooplankton) dominated the food base and these foods were taken 01' all feeding groups includirJg 0entbic feeders. Almost tunistic
summa rises
dll
species
employed
oppor-
feeding strategies. Table 4 the biomass, composition ared of this surf zone fish
food consumption
assemLJlage.
to no to
abundance,
biomass occur
and around
diversity
often
consecutive variable.
~~estill know very little of surf zone icbtbyofauna and the danger of sampling in heavy surf will proLably keep the
She
zone fish assemblage was a highly variable community whose structure and were probably dominated by dynamics factors while biological abiotic interactions were of secondary importance. As most food organisms were highly motile, their availability would be strongly influenced by surf conditions which in turn are determined by 'wind effects on the wave regime. Hence short term variability is largely
To really resource partitioning and niche structure of surf zone fishes some
situation understand
that
\Jay.
more accurate means of sampling is necessary whereby tbe actual location of fish within can lJe made. the
could spatial three
uifferent perhaps
In of
parts
SCUBA
of the surf
counts are
the distriin in the terms zone surf of in
solution.
and lJe better
this way
fish understood of
bution
feeding
partitioning dimensions.
fish
this
However,
\"ill lJe more
zooplankton
important
as food for
349 Table 4. Biomass, composition and food consumption of fishes in less exposed surf (After Lasiak, 1982). zones in the East Cape. FISH Number of Species Feeding group Biomass g.m-l (dry mass) Dominants 78 60 10 10 158 3 4 2 2 Other 6 5 4 0 Species or group Macrobenthos
I-1y sids/pr awns
-B1
1798
at night and studies not employing after dark sampling will miss this. Surf zone ~ish communities with are few highly
there are few published reports on the ecology of birds on sandy beaches. Most papers simply record counts of waders and other birds along sandy shorelines. Fitch (1950) recorded gulls dropping
dynamic species.
assemblages
in space and
time and short term variations in wind, conditions, are controlling surf probably major factor controlling a Most species show a these communities.
11igh
them and Brunton (1978) predation toheroa of (1980) dead veitch recorded beaches records (Kuyken, 1978). in New Zealand and have Leen made in and 1978) Britain Prater, Davies (1978)
on
and
degree of opportunism in feeding, consequently all feeding types may heavily sChooling on zooplankton when and they crustaceans
predate larger
are locally abundant. Planktivores, and benthic feeders with opportunistic are therefore p1anktivore tendencies, a major component of surf zone
important are as Fish energy transformers in surf zones and, because they are highly motile, are also important in the export of energy from ichthyofauna.
sanderling overwintering on recorded beaches, British while Schneider, Harrington (1981) demons t ra tecJ depl et ion of benthic .i?reyat a migratory stopover Burger et al. (1977) for shorebirds. compared the utilization of an open beach, a sheltered ueach and a salt marsh several
ad j acent
by
migrating
to beaches,
birds.
~here
are
te rns
reports 1977;
breeding 1981;
on or
pa r ti cula r ly
(Fuchs,
Randall, (Summers,
7. BIRDS
brief mention of birds as predators on sandy Koepcke, 1952; beach benthos (Koepcke, Vlhile Brown, 1964; Leber, 1982; Vader, 1982) some authors have made
plovers
350
cooper
amounts
beach and
of
insects
cast
!Jlowing ashore.
onto
On
months which revealed eight migrant and seven resident species. Summers et al. (1977) reco rded higher bi rd numbe rs on the west coast than the south coast of the western cape, presumably due to greater food abundance in the former area. 99,6km They recorded 1267 birds over of sandy beacbes and 8319 bi rds shores. on the Twelve beaches
car r ion
the kelp
dominated
on the \ve st coast birds are the main predators and were to take estimated 40% of herbivore standing stock (Griffiths et al.,
1983 ). Hoc key eta 1. (l 9 8 3 ) s u mm a r i sed
beaches
on sandy l.Jeaches
with sanderling (Calidris alba), whitefronted sandplovers (Charadrius marginatus) and black oystercatchers (Haematopus Resident moquini) most important. 7,3km-l and species averaged
in Soutb Africa. Tbey found diversity and al.Jundanceof shore!Jirds to increase with latitude. Birds consumed 10-49% of production macro!Jenthic different in areas and return faeces, feathers
beach system.
significant and
amounts
of
carcases
to
the
migrants 15,2km-l in summer. Summers, cooper (1977) recorded fewer oystercatchers on beaches than rocky shores and Summers, Haltner (1979) recorded mass changes in waders, showing most species to be heavier in winter. On the Natal coast Joubert (1981) made counts
Moran, Fischelson (1971) recorded two plovers, Charadrius hiaticula and C. feeding alexandrius, mysid on the
Ga s t r 0 sac
plovers
over 10 months, recording 14 species and densities of 39km-l, with waders making up only O,52km-l. Terns
roosted on the beach numbers. In the Eastern Cape (1980) counted birds over
wit h
in relatively
high
where mysids were densest. However, (1 S81) reanalysed Schneider their data to show that c. hiaticula selectively takes larger sid s . Schneider (1982) studied the foraging behaviour of
lilY
(Arenaria
interpres)
feeding
on Donax variabilis on a florida beach and Pienkowski (1982) described diet and feeding of two vlovers on a sand flat in
Northumberland. night and
hey '.c' fed d u r i n 9 cla Y and
12
months,
9e
recording
a v era
n u m l.J r s e
1 8 , 9 k 1<1-1
which
waders
estimated
bird
accounted biomass
from four
They for 47%. and also food standard metabolic species which
volychaet worms and taceans dominated in the diet. et al. of in (1980) detailed ana lysed on
crus-
Myers
the l.Jeach
dominant
.
predation crustaceans
sanderlings
made up 95% of numbe rs From thi s they estimated that birds were taking 32% of available macrobenthic production from these beaches each year. This was mainly- in the form of sand mussels (Donax spp.), mussel siphons, and mysids. Birds also took significant
la!Joratory
bey showed how several ex per i men t s . 'i.' variables affect a sanderling's capture by prey Capture is determined rate. density, the sampling rate and sampl i ng
efficiency. penetrability
Sampling of tbe
rate
depends
on and
substrate
3S1 sampling size, inversely 10mm efficiency depth with prey depends and risk depth, on prey substrate varied very flow part
!Jetween
of
them. larger
penetrability. or shallower
Prey being
zoopl ank ton, fishes and !Ji rds, while the interstitisal fauna constitute food web in the sand (McIntyre, Murison, 1973; McLachlan, 1977 a; a1.,1981a). 'l'he only McLachlan et links !Jetween
60% of prey
taken and
much less below 10mm. Larger prey were more vulnerable than smaller prey but small prey were never rejected. Prey risk was higher where prey organisms occurred close together. Capture rate could be predicted on the basis of prey size, density and sand penetrability. Fo!:" prey it was best to be small, deep in the substratum and not near other individuals. Prey species had no effect (Excirolana vs. Emerita). However, this was all done with dead prey. Most feeding although ac wrack was Jdone wi thout visual cues sanderling may use visual cues e.g. when feeding on beach Although probe locat ion by
these two food \4ebs is the common input of particulate (and probably dissolved) organics to both and the possible washing
(l-1U
of micro-organisms
tal., 1 9 78 )
n roe
more sheltered In situations, however, deposi t feeders and Iuay take large sand swallowers in amounts et al. , amphipods of interstitial fauna. Oliver (1982) recorded phoxocephalid feeding
filter
feeders.
times,
insects.
on a var iety of foods including nematodes and Cro ker (1967b) recorded a baustorid umphipod feeding on meiofauna, both cases being in sheltered lagoon sands. On beach Alheit, a (lS82) juvenile Scheibel recorded demersal fish feeding copepods. as BOl-/eve r , are mostly negligiule on look separately at the on
open
Iliac rof
harpacticoid exchanges
are
impo
rtant
preda to r s
on
these
beaches,
being
efficient during
croppers of
beaches
auna
of
~ne
macrobenthos
periods
food
lov: :.ide. On beaches where high inputs concentrate macrofauna of macrodebris along the drift line above the reach of ~arine predators, bi rds may be the most impo rta:-, t predators. Even bi ras that may be or.ly beaches :oos:. the on important 'rdhe the faecal deposits of re large flocks may add significant of nutrients locally. amounts
Several
authors
have
Itlen t ioned
aspects
of 1Jredation or food chains sandy L>eaches (Pearse et Bonnet, 1946; Koepcke, HedSJ?eth, 1957; Brown, a1., 1970b; Philips, 1973a, 1973b; Reilly,
Koepcke,
8.
8.1 Introduction The ~acrofauna and interstitial open sandy beaches form two communities with little fauna of separate no energy
Ansell et al., 1978; Reilly, 1979; McLachlan et al., 1981a; Leber, 1982; Gianuca, 1983). Macrofauna food chains start and end in the sea, wi th the land playing only a negligible rele, mainly
i n t 11e t 0 rTi1 0 fin s e c t s b low i n g 0 n tot 11e
or
,,~') .:J:) -
moving onto the beach to feed. Open beaches are characterized by the absence (Brown, 1964) and of attached plants primary production by the benthic microflora is generally negligible. On temperate beach, however, a sheltered (1968) found viable Steele, Baird
Ansell two
(2)
et
':jroups
a1 ., of and
only
(1) and et
herbivores
separated
carnivores.
0.1., (19810.)
filter
feeders
and
scavengers/predators
diatoms mixed to 20cm depth in the sand primary production but low and They measurable. considered vertical mixing due to wave action to be the main the shallow limiting factor. In beaches, subtidal off two Venezuelan Edwards benthic (1973a) primary recorded production
significant
kelp
Leaches
Griff iths et
herbivores,
(lS83)
uistinguished
carnivores
possible of
filter
and sandy
feeders.
trophic
'l'he
oou
sources
':Jroups
on listed in ~able 5.
macrofauna
Geaches
are
due
to
both ~re extremely important in surf water, altbough largely ingored to uate, '.,ould fall in cate(4) . On TIIOst sandy (1) and sories beaches toocJ source (1) is 01' far the most important, with food sources (3) (4) Leing and next. On sheltered shores, and
0 f
macro-algae (2943 and 6500 beaches) two but at the se areas are hardly typical of open beaches and the latter area VIas subject primary production to heavy pollution.
zooplankton,
yvhi
ch
may very column be water the in important, particularly where surf phytoplankton blooms occur. Unfortuna::.ely primary
however,
impo
cJetritus
production bloom
a surf
be
phytoplanktor.
for to
living
Til s t 0
filicroflora
may
oe
primary
estimated
prodllction
column
as
in the sediments, and unuer such conditions deposit feeders sucl1 as Arenicola,
incorporatec1
Callianassa
rtance
and
Scolelepis
may
be
abundant.
Different sources
C:: v'i"a:ls.
Hhile
Le<1ches
inputs
Lave are
or particulate
yet to
lJe
Til at
to some
JdeaSUreu,
distinguished
an
estiTi-lates requirements
available
Lased
TLiacrofauna.
on
the In
supply and a erratic of macrodebris supply of particulates while regular Hayes (1974) recognised dissolved and organics, stranded microparticulate plankton and large organic masses. Edwards (1973a) distinguished three main pathways by. which energy travels from primary foods to fishes; (1) via zooplankton to plankton feeding fish, (2) via benthos to demersal fishes and (3) algae to benthic directly from
of
the
India particulate 56141 kJ.m-2y-l 4291 kJ.m. -2y-l and 38765 m-ly-l at tively.
accounted consumption
feeCiers accounted for for and carnivores beach on Shertallai 235kJ. respecthus
1978).
the food Sl% of ., on average (Ansell et 0.1 In tbe L;ast Cape, filter feeding were estimated to assimilate
macrofauna
353 Table 5. Food sources and trophic groups of macrofauna on open sandy beaches. FOOD SOURCE 1. 2. 3. particulate organics including phytoplankton Deposited detritus, benthic diatoms, interstitial fauna FEEDING GROUP Filter feeders Deposit feeders
Scavengers
herbivores carnivores
4.
Predators
109421
kJ.m-.l.y-.L,
93%
of
the
macrofauna,
and fly larvae (Muir, 1977; Koop, Field, 1980, Stenton-Dozey, 1981; Griffiths, 1980; Griffiths, Stenton-Dozey, 1981 ; Koop et al., lS82a; Griffiths et al. , 1983). Biodegradion of macrophytes by the macrofauna In sheltered amounts of
deposited: high
making
up
remai ning 7% (McLachlan et al., 19 81a) . / This does not include zooplankton organisms feeders and which very are
abundant.
are only important where there are large inputs of debris or where filter feeders physical conditions or the absence of food (e.g. East Coast of South Africa, Dye et al.,
a reabsen t becau se of
unstable
localities or where large particulate organics are deposit feeders rllay reach
(Eleftheriou, values McDermott, 1983) On 1976; most beaches organic content of the sand
lJiomass
MCIntyre,
is beaches adjacent to kelp beds or similar areas. In southern California Hayes (1974) estimated these inputs to amount to 473kg wet (ca.78kg dry) algae -1 -1 m.y (Macrocystis), while on \Jest Cape beaches adjacent to kelp beds
very
low,
e.g.
0,2-0,4mg.g-l
sand
in Scotland and India (Steele, 1976) and undetectable amounts in the \Jest cape and Bast cape (BrOvlD , 1971a; McLachl2.n, 1977a)
.
Predators on the other benthos include crabs, starfish and gastropods. These normally abundance, occur making in up relatively 5-10% of low biomass.
wet Ecklonia
KOOp et Griffiths,
1980;
Griffiths et aI, 1983). While (1974) found Tylos punctatus to only 4-5% of the for estimates taken by edible kelp macrofauna amphipods
As many are supralittoral forms, e.g. Ocypode, they may be the only macrofauna where condi tions are too dynamic macrofauna to survive for intertidal (Gauld, Buchanan, 1956; Dye et al., 1981). swash
consumption being
354 Energy flows through different species These food chains end in fishes, birds, crabs and other invertebrate predators very man. iew studies and have estimated different juvenile ant the flatfish importance are the of these predators. cropping Scotland In most importsiphons of
and trophic efficiencies vary widely. Turnover (P/B) ranges from 0.3 for tenuis on a temperate beac h Tellina (Trevallion, spiculum on 1971) a to 10.3 for Donax beach tropical Indian
(Ansell et a1., 1972b). Ansell et a1., (1978) found turnover on these tropical beaches generally ten times higher than on cold temperate Scottish beaches. Edwards (1973a) recorded average P/B's macrobenthos, of 2.8-3.5 for the t-1cLachlanet al., 0.5-5.0 molluscs for and (1981a) used P/B's of production of Koop, crustaceans, and the P/B for Ligia somatic
predators,
bivalves, palps of polychaetes and whole prey. Bival ves may then put more ene rgy into regeneration of siphons and less into reproduction. Trevallion (1971) up to 1.3% of total estimatec.1 that energy annual expenditure in 'J.'ellina may go into regenerating tenuis siphons. the most In India crabs were considered
important predators, with fishes and man less impor tan t and birds virtually absent (Steele, 1976; Ansell
dilatata at 3.75 times per year. Assimilation 100%) vary efficiencies widely, commonly (A/C being X very
low for scavengers/grazers feeding on abundant macroalgae, e.g. 64% for 'rylos (Hayes, 1974), 5.5% for Ligia (Koop, Field, 1981) and 30-50% for Talorchestia and fly larvae, (Stenton-Dozey, 1980). Koop et a1., (1982a) average overall estimated an efficiency of for assimilation 22% invertebrate kelp grazers while Ansell et a1., (1978) and Edwards (1973a) assumed assimilation efficiencies of 80% Griffiths,
on the other beach. McLachlan et predation al. , (lS81a) partitioned on east cape Leaches. Birds took 32% of intertidal macrofauna production al., 1980), fishes were (t-1cLachlan et estimated predators to take 55% and Genthic (mainly crabs, Ovalipes) 10%
13%
with Elan accounting for less than 3%. DU Preez (1981) has subsequently made a detailed of
La
for
carnivores
investigation
of the predation
Ovalipes
s i a k
fish
beaches
on Donax and Bullia and and Rossouw (1983) have studied predation. kelp On dominated
of the Hest Cape, 110weve r, bi rds (Griffiths
in India while Edwards (1973a) estimated p/ 17% C X 100% as 15% for benthos, for demersal fish and 11-15% for other fish at his unpolluted site. Ecological efficiencies calculated from et a1. (1981a) were 9-19% macro-fauna with a mean of 17%. McLachlan for the
predators
et
Evaluating our knowledge of sandy l>each food chains as a whole is made difficult by only not the small number of quantitative studies, but also the absence of data on zooplankton. The
355 macrofauna cannot be considered in isodescribing these food lation when chains. production, Surf zone also untimately zooplankton falling prey To conclude this section on macrofauna energetics it appears that five main beach types can be recognised in terms of macrofauna energy flow patterns (Table 7). Information on surf zone zooplankton in different zoogeographic regions is unfortunately not available to allow incorporation into this scheme. It may Le expected, however, that as zoopolankton and macrobenthic filter feeders utilize the same food resources, their
similar
to fishes and being exported from the system, is probably more important in terms of energy for example, surf flow. In the East Cape where we have some idea of
zooplankton biomass, zone a pattern emerges if larger different mobile crustaceans, such as mysids and prawns, are included with zooplankton
total
biomass
vv'Ould Le
roughly
out to 500m from the beach (Table 6). Table 6. Summary of approximate faunal dry biomass for an average beach and Cape. surf zone East (From in the McLachlan, 1983) BIOMASS SJ .m -1
in filanycases.
been
paid
to
CA7E;GORY
!'Jacrobenthos
Zooplankton
Birds Fishes
materials in the sand. Little was made subsequently to evaluate or quantify this until quite recently. system This consists typically of bacteria, protozoans and organic attempt meiofauna constituting their own tood web in the sand.
The food chain is centred around filter feeding feeding benthos and zooplankton particulate detritus and phytoon plankton and being taken in turn by fishes, birds and crabs. As zooplankton turnover is 6-7 times tbat of the macrofauna, the centre of gravity of the food chain is clearly in the surf and not on the beach. HcFar land (1963) came to a similar conclusion, showing that plankton was the predominant fish food the The off Texas beaches, greatly exceeding benthos as an available food source.
Dissolved and fifle }?articulate organics form the Lase chain, of this food although in sheltered situations the benthic microflora might also play a role. McIntyre et al., (1970) were the first to demonstrate that interstitial
fauna '2.)uld subsist on uissolved organics. Using two experimental sand columns, they maintained populations viable microand meiofaunal in the second
column on the effluent leaving the first column. They suggested that bacteria were the prime utilizers of uissolved
organics, utilizing
these
org ani c
mo le-
356 Table 7. Major beach types in terms of macrofauna energetics. Predominant Sources Food Dominant Trophic Group Filter Feeders
I-1 c r 0 f au a
n a 1
Latitude Tropical
Predators
Particulates
Invertebrates Fishes
Temperate
Particulates
Low to high
Filter Feeders
Fishes,
Birds
Invertebrates
3 .
r.1 c rod a
e !J r i s
based
food
or Tropical
Low to high
Deposit Feeders
Birds,
Fishes
Invertebrates
Temperate or Tropical
S t ran
d 2
(I
c 2. r r
ion
:",0'(/
beaches. where they can be digested with extracellular enzymes. The meiofauna in turn fed mainly on these bacteria, accountinCj for approximately 5% or bacterial consumption.
some
directly dissolved
1.1 i 0 f au e
n a 1
species
may
!Jroad
feed
meiofauna
on
organics,
Fenchel (1972) considered bacteria the only primary decomposers of particulate plant detritus with their decomposition rate limited by Consequently he trophic levels oxygen and nutrients. suggested that higher food
stitial food we!J as most species feed on bacteria, protozoa or other meiofauna (McIntyre, Murison, 1973; McLachlan et 1981a). ale , The position of protozoans in this system has !Jeen little studied in open beaches; \/h i 1 e some forms may have a similar trophic to status are even predatory on meiofauna. However, in a broad sense they probably occupy a .tJosition interbacteria,
in the interstitial
others
357 mediate between, and overlapping with, bacteria and meiofauna in interstitial food chains. Three within herbivores trophic the and types may be
recognised predators, with only
has been the stuuy of experimental columns in the laboratory. Several authors
meiofauna, bacterivores,
have maintained laboratory sand microcosms for extended periods (McIntyre et al., 1970; pugh, 1976; Boucher, et al., 1977; McLachlan Chamroux, Hormald,
19S1c)
the Hydrozoa and Turbellaria exclusively predatory (Tietjen, 1980). Swedmark (1964) however, listed four categories, predators, diatom and epig rowth feeder s, detritus Besides the side red eaters direct that and suspension of feeders. on congrazing meiofauna (1980)
1976;
~tirling,
.
Chamrou x
microfauna,
Tietjen meiofauna
stimulate
al. , circuit system has been maintained for 16 months on soluble amino acids (Boucher, Chamroux, 1976) confirming the utilization of dissolved by microorganisms organics and their subsequent However, structure indicated soluble
the In only this
et
bacterial activity in the interstitial system by (1) mechanical breakdown of det:itus, bacterial nutrients secretion bacterial conveying making for c010nisation, to it more susceptible (2) excretion of (3) use, microbial
(decreased
!Jacteria
utilization was
the
of mucous thereby attracting (4) growth and movement nutrients and oxygen. efficiencies (A/ X C have been estimated
llIeiofauna.
!Jacteria
meiofauna
Assimilation
to account for 95% and 5% of input respectively, the same y r e as f 0 u n d by 1.1 In t c eta 1 .
in the range 6-26% while transfer efficiencies (P/ X 100%) were A for three nematode species 79-97% (Tietjen, 1980). For turnover (p/) B a ratio of 10 has been widely used for meiofauna
Because 'working of
(Gerlach, 1971).
the d iff icul ty minute of scale when of
with
the
inhabitants
w'arrant attention concern a comparative study of Scottish and Indian beaches, studies on high energy beaches in the Cape Eastern and studies on beaches
...
fauna, investigations the interstitial energetics interstitial have of generally taken the form of 'black box I studies has of tbe systertlas a \Jhole. This been or done by oxygen measuring consumpbenthic commonly
receiving
high
kelp
inputs
in
the
al.
"(1978)
Illade
a
of
detailed
investigation on
dynamics
tropical
interIndian
(Md 150-190u!l1) and Scottish (Md 250um) using small and large
tion measurements have been discussed and reviewed by Lassere (1976), Pamatmat
35'S
diatoms which were common on the temperate beach. Dissolved organics were estimated to make up 80% of the input on the Scottish beach and 39% on the Indian beaches. Interstitial community respir(164gC.
vJinter
India. I-loweve r , surf flushing of the Indian ueaches way remove significant
quantities of bacteria, thus making them available to macrofauna filter feeders and decreasing the quanti ty availal:Jle to
the meiofauna.
ation on the tropical beach m-2 .y-l) was nine times the values and twice the summer the temperate beach (mean y-l) with microbial estimated to be an 72 and former 15g.c.m-2.y-l
gc.m-2.y-l
average annual of (using the latter in respiration figures based on an assumed conversion efficiency of 45% and the assumption that the contribution of Meiofauna meiofauna was negligible). biomass was, however, much higher on the Scottish the 8) . beach (273-523mg. m-2) than Indian beach They postulated (24-60mg.m-2)(Table that more vigorous
In the Last Cape the interstitial fauna consists essentially of meiofauna, protozoans and bacteria to considerable depth in the sand (Dye, 1979a; NcLachlan et al., 1979a).
f 1 uc t ua t ions in
tion as a result vf tidally induced changes in pore moisture and interstitial water flow, lowest values being recorded when the sand dried out. Dye (1979a) investigated biological oxygen demand in sand cores in as these gave higher the laboratory values than in situ techniques. He showed that caused disturbance a significant increase in oxygen uptake. study Dye (1981) more cJetailed partitioned this oxygen uj?take l:Jetween bacteria, protozoans and meiofauna on two beaches (TaLle 9). In a later beach sand, using intact
flushing of the sand on the tropical Indian beach both boosted respiration bacteria from the sand and striped grains. tropical beach received more The organics and had a higher interstitial consumption rate than the Scottish beach as a result of increased water flow by wave flushing. Surt oxygen
~able 8. Biomass, production and respiration of interstitial organisms on Scottish and Indian beaches. (After Munro et al., 1978). All values in gc.m-2.y-l. INDIA Production 0 72 2 SCOTLAND Biomass
0 31 0.02 4
Respiration
Production
7 15 2
Biomass
0 160 4
0.2
359 Partitioning of interstitial oxygen uptake on two East Cape beaches. (After Dye, 1981).
MODERA'l'ELY EXPOSED BEACH
Table 9.
Biomass %
02 uptake %
75 19 6
18 18 64
79 17 4
21 24 55
In the West Cape very rich interstitiul populations develop on beaches wi th high Koop, Griffiths (1982) kelp inputs. recorded dry biomass of 241, 200 and 663 g.m-l of macro- meio- and microfauna respectively. Bacteria were concentrated at lower tide levels, probably subsisting on small particles of broken down
fa.eces Koop
rap i d 1 Y .
1:' beg
rea t
s i g n i f i can ceo
these (1974)
microbes recorded in
Hayes kelp
breakdown
he
kelp of
8.4 Conclusions It may be concluded that intertidal sandy beaches are important in the processing ot organics whether these be as DON, from POH or larger animal and plant toods genere.l1y remains. These COllie
flow much
for of
the kelp resulted in high concentrations of leachates (up to 5640 mgC.l-l) in the sediment below the kelp. Over 90% of this was utilized by bacteria after seeping through 1m of sand and 23-27% of carbon in the kelp was converted to bacterial carbon. The ratio of increase
the sea and in turn most production returns there. The interstitial fauna alv;ay s more important are than the macrofauna of in this mineralisation organic materials, even wbere a rich macrofauna occurs (Table 10). The
interstitial
system
is
Criven
by
in bacterial biomass to kelp carbon used giving conversion a was 58. 7% , efficiency of 29.4% if bacterial biomass is
~1u h c
50% carbon
of the
soluble and !?articulate being via the main }?ath bacteria attached to sand grains to meiofauna. The intermediate rale of protozoans is uncertain. 0n exposed
organics,
input s
of
grazers were also converted to bacterial grazers these have low carbon as assimilation
remaining mineralised
beaches there is little excbange between this sytem and the mac rofauna although the importance of microorganisms flushed from the sand as food for the lilacrofauna needs investigation. further The concentrations of meiofauna ilushed from
360
Table 10. Approximate partitioning of total benthic assimilation between filacrofauna and interstitial fauna in four geographic areas based on data in Munro et al. (1978) (1982). Ansell et al. (1978), McLachlan et al. (1981a) and KOOlY, Griffiths,
Scotland, temperate
30 000
(23 )
S9 000
(77)
60
India,
tropical
427 000
(38)
703 000
(62)
60
116 000
(37)
236 000
(63)
GO
42 000
(3)
1599 000
(97)
63
exposed beaches here sand of the measu~ed by McLachlan (1977a) and found On sheltered beaches to be negligible. a greater size spectrum of organisms may occur, filling the gap between interstitial fauna and larger macrofauna and thereby facilitating energy flow by predation. In such environments deposit feeders may also ingest significant quantities of interstitial fauna. Inshore ments zone food are chains in beach environhowever centred in the surf than the intertidal, even
9. may
NU'I'RIE;N'l'
CYCLING
been specula ted tha t beaches in recycling importance
It has
long
(jf
Le
organic mineralising materials coming from the sea (Pearse et al., 1942) . This is effected Ly the nutrients
Ly
orSjanic anu in
nutrients
some areas PO4. In NH4' groundwater seepage can, bo\vever, also supply considerable amounts of inorganic nutrient via the Leach to the surf zone. This can take the form of Goth artesian aquifers ariel 'jeneral seepage around the
area
rather
where rich intertidal macrofaunas occur. Our area of least understanding at the moment is these dynamic food chains of the surf zone and in particular the role zooplankton and larger motile of
crustaceans.
meets this
the permanent water taLle the sea. Johannes (lS80) fauna to Le very important in Hestern where
\,her e
Australia
submarine
grounell,Jater
361 discharge ranged 1-5m.y-l and connitrogen (Chamroux, 1965; McIntyre et al., 1970; pugh, 1976; Boucher, Chamroux, Stirling, McLachlan 1976; 1979; et is Cahet, Munro the 1976; Wormald, et al., 1978; Gener ally activity
tained up to 380 ug at 1-1 N03-N. Nutrient regeneration by the macrofauna, zooplankton, birds and fishes has been virtually (1979b) and respectively excretion in Siliqua and Washington and ignored. Lewin et al., (1983) ammonia Prosch, McLachlan investigated
al. ,
19 SIc) . dominant
nitrification
in Leach sand and organic nitrogen is efficiently mineralised to nitrate. Increasing organic loading tends to raise equilibrium levels and it is usually only at very high levels that a beach can not cope and anoxic conditions and ammonia appear (Oliff et al., 1970; pugh, 1976). Boucher, Chamroux (1976) recorded total Itlineralisation of amino Munro acids adcJed to tbeir sand columns while et recorded al., (1978) 70%
Africa. In both cases very large populations of these bivalves develop and nut rients regene rated by them could supplying significance in be of phytoplankton requirements in the surf Hayes Tylos to (1974) found zone. only on 4-5% of kelp input beaches while Griffiths, Californian Stenton-Dozey (1981) and Koop et al. consume (1982a) estimated kelp was that 9-74% of stranded by the macrofauna. consumed
~1cLachlan et mineralisation
65% when amino these
increased
to
Not all of this is assimilated, however! may efficiencies and low assimilation comparatively a small result in being kelp nitrogen prop6rtion of regenerated directly by t.he macrofauna. The contribution of other faunal components must also be important in this regard but have yet to be investigated.
Generally .3andy
important
acids
acJded.
of
.studies
somewhat been
water situ
unrelatecJ
there has been continuous submerand inputs have been constant than tidal or pulsing. Except-
the
in
interstitial
has been
system
of
more
beaches
thought
nutrient
regeneration
the
macrofauna.
Water
filters
than through
ions are pugh (1976) and McLachlan et al. (lS81c), although even these experiments Jid not incorporate pulsing flows. Filtration rates have ranSjed 25-1000 1.m-2.cJ-l, covering a variety of flows both intertidal and subtidal.
t.lcLachlan
the sand, as a result of tides and waves both in the intertidal and the subtidal, and DOM and P 0l'1 are removed Ly tbe interstitial fauna and mineralised. Several studies of experimental systems
earlier sand col umn \vor k with a model of water filtration and nutrient regeneration by sandy ueaches. This was based on a regression model a predicting of ancJ filtration particle range. tide as function slope sand
(1982)
followed
up
of sand columns or model beaches have looked at mineralisation and/or nutrient mostly concentrating on regeneration,
volume size,
beach
of the in
of mineralization
of organics
to rocks and subject to larye kelp inputs are interesting for comparison. It may therefore be concluded that sandy beaches can mineralize most oryanics tl1ey receive, the l>ulk of this l>e ins
done very efticiently fauna. (80-100%) This l>y the interstitial
relation to sand particle size and the average distance. The filtration filtration distance is about 35% of the (Riedl, distance intertidal horizontal Machan, a 1972) and average mineralisation At rate of lO41.m-l.d-l was estimated to be at least 60-80%. filtration
is water
accomplished
uy
the l?rocess of
and an organic nitrogen level of 0.15mg. 1-1 this predicted regeneration of 0.9g N03-N m-l.d-l. An attempt to by by estimate total beaches in the nutrient production East Cape was made
filtration through the ueach and will be greatest where steep beaches and short period waves result in high filtration volumes (e.g. in the tropics) . Hhere vast macrofauna populations occur tiley
also
contrilJute
They estimated
may
significantly
to
nitrogen excretion by the macrofauna and interstitial fauna of the beach and surf 350gN.m-l.y-l 591gN. at and zone -1 .y-1 respectively. on Thus, even m as these with very high beaches such biomass, the interstitial macrofauna fauna are more important in nutrient regeneration, accounting for 63%. Hhile figures approximate, the above only suggest that the mineralising activities the of the benthos alone can replenish nutrient pool of the surf zone in a matter of days or weeks. Koop bial et al. (1982b) quantified micro-
regeneration.
Zooplankton,
iishes
must
also
ue
very
1'-10st organic
nitrogen
0.
and will
phosphorous rapidly
supplieCl
to
IJeach
ue
to returned the surf as nitrates, ammonia and phosphates. Thereafter the fate of these nutrients will depend on physical conch t ions in the surf 20 ne . Hhere cellular circulation patterns occur there may Ge a high degree of retention, coasts or reflective whi Ie on irregular
beaches,
exchange
regeneration of nitrogen from stranded kelp over an eight day cycle. In this time the microbes incorporated
Ge much more rapid. As I.lany surf zones are characterized l>y high primary productivity, utilization of Geach generated nutrients in the surf 2.one way be very important and is in need vf more Hayes (1974) and Koop et al. attention. (1982b) consider such Ilutrients of impurtance little in the surf Vlbile McLachlan et al. (1981a) suggest that these nutrients may l>e sufficient to surf phytoplankton Glooms on maintain long open beaches. 10. This
TmJARDS A l-10RE;
94% of the kelp nitrogen, which normally strands at an average input of l2gN.m-l.d-l. However, in the long term the beach can not act as a nitrogen sink and this must return to the sea. In this case this could supply roughly one quarter of the nitrogen of phytoplankton in the requirements kelp adjacent
beds. Hhile hardly typical of open beaches, even on the west coast (Bally, 1981), these figures for beaches adjacent
revieVl
has
363
look at our present knowledge of the ecology of beach/ surf zone env ironment s as a whole, fauna and incorporating flora, all aspects of flow and energy
Energy flovl and nutrient cycling in a long open lJeach typical of the Last Cape is illustrated in rig. s. It This is a of rich system energy and producers (benthos, decomposers Energy flows to be boundaries.
\v'i tb
considerable
flows
have not attempted such a holistic approach and have tended to investigate one or more components in isolation. In the East Cape research on beach
(interstitial fauna). within this system appear than across its However, although exchange
greater
has been unique in holistic approach towards quantification of energy flow and cycling zone under the concept This body of a is
ecosystem.
across the dune/beach boundary is negligible, exchanges across the outer boundary
quantified.
beach/surf
(HcLachlan,
from tine organic materials (POM phytoplankton) via L.ooplankton filter-feedin';j Ti\acrofauna to The interstitial
important
and and
beach
together
the water
of the envelope
of the surf zone form an ecosystem where internal energy flows are greater than exchanges across boundaries and (2) that boundaries, this system has definable these being
mar
in
mineralisin<;J
The
total
c.mount
lJy the
regenerated
surf
60 ne
fauna is not yet known, Lut must De SUDar,d interas the Iliacr of auna stitial iauna alone regenerate can enough to replenish the zone surf nutrient pool within a few weeks to days. Altbouyh not indicated in Fig. 9,
1 and via r d
stantial
surf circulation cells on the seaward margin. The central concept is that the circulation cells that typify the surf zones 1973) of open beaches (Inman, Brush, imply a certain measure of
wave action probably represents tilemost important factor controlling this system, as it is responsible for transporting water, cells sand. not only biological ancl needed to information is understand beach/surf ecosysteIlls. Ijater chemical movement, processes the. key
occurring
integrity retention
or
surf
zone
water within
and this
zone. that
materials
materials,
sediment biological and driving surf circulation and pumping water through Leach
between sea interfaces and land. Clearly, however, this can not apply to poc ket beaches, lJeaches assoc iated with
Clearly,
rocky shores or beaches dropping directly into deep water where there is
no real .surf zone. The kelp input beaches of the South African west coast, example, as interfaces function for between adjacent kelp beds and the land.
to here,
surf
oceanographer.
exchanges
across
them
364
SEA
I
SURF
EXPORT
BEACH
OTHER PREDA TERS 1837
3684
DUNES
8861
I
I 2468
45:70
IMPORT 8337 600
1
EXPORT 2155
I I I I
.
=
I I
:
I I
I
.
I
I i
t I
!
I !
8-t I
I I
236325
:612~
. 89269
I I I \ I \1 ~ I I ... i I
I I I I i I I I
J ~
LINE
DRIFT
FIGURE 9. Energy flow (solid lines) and nutrient cycles (broken lines) for a typical East Cape beach, largely after McLachlan et al. (1981a). All values in kJ.m-l.y-l except nutrient values in g inorganic N. need to be quantified. essential that physical biologists Beaches need
study
It is therefore oceanographers, work this together. areas only must fashion. geographic
ecosystems,
evaluated
knowledge environment.
and this only can be in the presence of a detailed of the physical and chemical
and be
chemists in
in
different studied
to
the
Biologists
REFERENCES Achutankutty CT (1976) Ecology of sandy beach at Sancoale, Goa: Part 1. physical factors influencing production of macrofauna, Indian J. mar. Sci. 5, 91-97. Alikunhi I( H ( 1 9 4 4 ) '1'he z 0 n a 1 d i s t r i bution of the mole crab (Emerita
venture out into the surf zone. In the dynamics organic of fine materials, phytoplankton and zooplankton lies the key to a more realistic understanding of the function of sandy 0each/surf zone
365 asiatica) on the 11adras coast, J. Bombay Nat. Hist. Soc. 45,94-96. Alheit J (1978) Distribution of the polychaete genus Nephtys: a stratified random sampling survey, Kieler Meeresforsch 4, 61-67. (1976) Alheit J and Naylor E zonation Behavioral basis of intertidal exp.mar. in Eurydice pulchra Leach, J. BioI. Ecol. 23, 135-144. Alheit J and Scheibel W (1982) Benthic harpacticoids as a food source for fish.
l-1a r. B i 01. 70
the
macrofauna,
Mar.
Biol.
17,
38-62.
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