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Thirty years of monitoring traditional fish trap

catches at Kosi Bay, KwaZulu-Natal, South Africa, and
management implications
a
R Kyle
a
Ezemvelo KwaZulu-Natal Wildlife , PO Box 43, KwaNgwanase , 3973 , South Africa
Published online: 09 May 2013.

To cite this article: R Kyle (2013) Thirty years of monitoring traditional fish trap catches at Kosi Bay, KwaZulu-Natal, South
Africa, and management implications, African Journal of Marine Science, 35:1, 67-78, DOI: 10.2989/1814232X.2013.769905

To link to this article: http://dx.doi.org/10.2989/1814232X.2013.769905

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 African Journal of Marine Science 2013, 35(1): 67–78 Copyright © NISC (Pty) Ltd
Printed in South Africa — All rights reserved AFRICAN JOURNAL OF
MARINE SCIENCE
ISSN 1814-232X EISSN 1814-2338
http://dx.doi.org/10.2989/1814232X.2013.769905

Thirty years of monitoring traditional fish trap catches at Kosi Bay,

KwaZulu-Natal, South Africa, and management implications
R Kyle

Ezemvelo KwaZulu-Natal Wildlife, PO Box 43, KwaNgwanase 3973, South Africa

e-mail: rkyle@iafrica.com

The catches of the traditional fish traps in the Kosi Bay estuarine lakes were monitored over a 30-year period from
1981 to 2010. Monitoring data were used to provide estimates of, and insights into, catch size and species composi-
tion as well as seasonal and annual cycles of catch abundance. Over 1.2 million fish, comprising 43 species and 23
Downloaded by [University of Saskatchewan Library] at 04:06 28 December 2014

families, were estimated to have been caught during the study period. Tag-and-release data were used to estimate
the impact of trap fishing on fish stocks and comparisons were made with recreational fishing to indicate overall
fishing pressure and the sustainability of the fishing. Trap numbers remained fairly similar from the first year of
monitoring (1981) until 1994, but thereafter they more than trebled by 2001; although numbers decreased after this,
they remained well above earlier levels. During this period, recreational angling was an important factor, and gillnet-
ting — both legal and illegal — added to the fishing pressure. Information from the fish trap monitoring, together
with results from fish mark and recapture studies, suggest a very high, and possibly unsustainable, catch rate that
requires management intervention to return them back to historical and sustainable levels.

Keywords: catch monitoring, estuarine, fisheries, fish marking, fish traps, iSimangaliso, overfishing, South-East Africa, World Heritage Site

Introduction

Semi-permanent traps are used in many shallow marine, Study area

freshwater and estuarine areas of the world to catch fish and
invertebrates (Mahon and Drayton 1990, Shaul and Reifsteck
1991, Beliaef et al. 1992, Russ and Acala 1996, Mwatha and
Orembo 1998, Siddeek et al. 1999, Copes 2000, Koranteng
et al. 2000). Some of these are traditional practices that
have remained fundamentally unchanged for many genera-
tions (Mwatha and Oremba 1998, Copes 2000, Koranteng et
al. 2000). Despite this, little work has been carried out on the
traps, their catches and impact on the stocks of target popula-
tions. Traditional fish traps have been in operation in South
Africa in the Kosi lakes for many generations (Felgate 1965).
They have been described (Tinley 1964) but no comprehen-
sive long-term account is available of their catch, either in terms
of species composition, overall size or their impact on stocks.
This study records the Kosi Bay traditional fish trap catches
from 1981 to 2010, and represents one of the first long-term
datasets for fish traps in an estuarine lake system. It was
initiated because of the perception of environmental managers
of the area, Ezemvelo KwaZulu-Natal Wildlife (Ezemvelo),
that there was conflict between traditional, rural, subsistence
trappers and the increasing numbers of relatively affluent
recreational anglers. During the study, the conservation status
of the area has been enhanced several times but, due to the
initial and apparently sustainable nature of the trap catches
(Kyle 1986), fish trapping was allowed to continue unaltered.
Monitoring of trap catches not only produced a long-term
dataset but also, through an associated tag and recovery
experiment (Kyle and Robertson 1997), provided important
insights into the impact of trap catch on fish stocks.
Situated on the east coast of South Africa, just south of
the Mozambique border, the Kosi Bay lakes are a series of
water bodies about 10 km long, running parallel to the Indian
Ocean immediately behind high vegetated coastal dunes
(Figure 1). In the north, the system opens to the ocean
through a permanently open mouth situated at 26°53′44″ S,
32°52′49″ E. From seawater conditions in the mouth of the
system, the lakes become progressively more fresh water
upstream; lake Amanzamnyama being completely fresh
water as it is not affected by tidal action. The lakes are fed
by two small streams, the Nswamanzi in the west and the
Siyadla in the south. Various aspects of the system and
its fisheries have been described (Tinley 1964, Hill 1969,
Begg 1978, 1980, Blaber 1978, 1981, Cyrus 1980, Kyle
1981a, 1981b, 1986, 1996, Cyrus and Blaber 1984a, 1984b,
Mountain 1990, James et al. 2001). Most of the fish species
of importance to humans are either estuarine or of marine
origin (Blaber 1981, Kyle 1986, Fennessy et al. 2001). The
lakes were proclaimed a nature reserve in 1988, listed as
a wetland of international importance under the Ramsar
Convention in 1991 and became part of the iSimangaliso
World Heritage Site in 1999.
Fish traps presently occur from just inside the Kosi
mouth into lakes Makhawulani, Mpungwini and Nhlange,
though there were none operational in Lake Nhlange
during the study until 2010. (Figure 1). The Kosi Bay traps
are semi-permanent structures, made from branches and
other plant material that are collected in the area around
the lakes. They are described in detail by Kyle (1981a)

African Journal of Marine Science is co-published by NISC (Pty) Ltd and Taylor & Francis
 68 Kyle

fish to the bank. In the past, all the materials used were

MOZAMBIQUE
collected from the surrounding indigenous bush; however,
in recent years, people have begun using gumpoles, nylon
AFRICA rope, gillnet and other modern materials. A 30 m wide
SOUTH KZN channel, from the lakes to the ocean, is kept clear of traps.
AFRICA Historically, the binding material used in trap construction
South
Africa
INDIAN was generally dried leaves of the wild banana Strelitzia
OCEAN nicolae and as these leaves are quite thick, there was a
Lake substantial gap between two neighbouring sticks, allowing
0 2.5 5.0 km Zilonde fish narrower than about 3 cm to escape. As the trap fences
Kosi Mouth curved upstream from the banks, they tended to catch
primarily fish moving out from the lakes towards the ocean.
The local residents who carry out the fishing are members
COASTAL FOREST RESERVE BOUNDARY
of the Tembe/Thonga tribe of southern Mozambique and
northern KwaZulu-Natal (KZN) in South Africa (Felgate
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Lake 1965). They live around the lakes in individual homesteads

SOUTH AFRICA Makhawulani and have access to the fish traps on a daily basis. The region
KWAZULU-NATAL Lake still has strong traditional tribal structures although elected
Mpungwini
officials and central government influence is increasing.
Kosi Bay The surrounding area is very rural, with little formal employ-
Camp Site
ment, and the community is heavily dependent on natural
zi
an

26°58ƍ S resources for their basic needs of food, shelter, medicines

m

and warmth.
wa
Ns

Lake Nhlange
Material and methods

KwaNgwanase The consultation process and preparation for monitoring

(Manguzi) Banga As a precursor to trap catch monitoring, the concept of
Nek
numbering and monitoring the traps was carefully explained
to the local Induna (Headman) and Nkosi (Chief) in order
to obtain their support to conduct the monitoring. This
Lake formal approval had the benefit of publicising the scheme,
Lake Amanzimnyama
Shengeza which was essential if complete catch monitoring was
to be achieved. It also ensured the compliance of all the
la
ad

trap owners, which may otherwise have been a problem

INDIAN
Siy

as measuring each fish caught would result in delays to

OCEAN
32°50ƍ E
Figure 1: Map of the Kosi Bay lakes, South Africa, showing
features important to the Kosi Bay traditional fish trap fishery
and Tinley (1964). In brief, two parallel rows of branches
are pushed into the sand about 50 cm apart. They usually
start at the bank and lead out into the channel, histori-
cally curving almost exclusively upstream. These are
packed with brushwood to form a fence to partially block
fish movement. At the apex of the fence is a heart-shaped
palisade to guide fish to a terminal basket with fish-proof
walls and a valve constructed in such a way as to allow
fish to enter but not leave. Fish, usually migrating from the
lakes to the ocean, travel through the lakes and channels
mostly at night and passively move into the ‘baskets’. In
the morning, the trap owner, usually a male living nearby,
spears the fish in the basket, threads them onto a vine
(or more recently a piece of fencing wire), and drags the
the trapper. Peer and community pressure was important
in ensuring the co-operation required for the monitoring
process. The local Induna provided candidates for
monitors from among the trap operators. These men were
interviewed and five of them who knew how to identify all
the fish species and could read and write were selected
and appointed. They were trained in the completion of daily
catch returns detailing the date, trap number, fish species,
number and total length of each fish caught. Monitors were
issued with metre-long measuring boards, datasheets,
pens and clipboards. Trap owners were generally found
to have a sound knowledge of most species of fish caught
and identifications were made based on Zulu names.
Problems were initially encountered with the identification of
some mullet species, but older, more experienced, fishers
were interviewed and Zulu names for all important species
were obtained. The only identification problems — which
remain unresolved — were the ‘lumping’ of Carangidae
and Sphyraenidae and some minor species (Table 1) as
local fishers considered these two families to represent
one species only and they would not easily accept other
identifications.
Monitoring methods
In order to record all catches, the traps were numbered,
 African Journal of Marine Science 2013, 35(1): 67–78
Table 1: The monitoring intensity and proportions of total catch
NUMBER OF TRAPS/BASKETS
recorded in the Kosi Bay fish trap catch monitoring programme
from March 1981 to February 2010
Number of Approximate
Period
monitors coverage (%)
March 1981–February 1985 5 100
March 1985–February 2000 1 12.5
March 2000–February 2010 2 30
mapped and then divided into five areas, the sizes of which
were such that they could easily be covered daily by a
monitor. Fish were recorded against the trap number but
most traps consisted of several fish-catching baskets. The
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number of working baskets in a trap could be increased or
decreased by the trapper at short notice and so the number
of traps is probably a better overall longer-term gauge to the
size of the fishery. Early each morning, monitors stationed
themselves at a site from which they could see all the traps
in their area. When a fisher left his trap he was intercepted
and details of his catch were recorded. Data sheets were
collected regularly and monitors were strictly supervised to
avoid circumstances (e.g. absenteeism) that could lead to
inaccurate results. From March 1981 until February 1985,
all traps were monitored (Table 1) and initial results were
reported by Kyle (1986). From March 1985 to February
2000, due to financial constraints, monitoring continued
in only one of the five areas, which had accounted for
approximately 12.5% of the total catch by numbers over
the intensive investigation period. In March 2000, another
monitor was added to increase the proportion of traps
covered. From 1985 to 2010, total catch estimates were
generated by extrapolation from the estimates of the
proportions monitored (Table 1). The first year of monitoring
(1981) consisted of data from March 1981 to February 1982
and subsequent years followed the same pattern.
Fish total lengths were analysed for the period 1 March
1983 to 30 April 1994 to give information on the length
distribution of the important species. Estimates of fish
mass were calculated from weight/length relationships
provided by van der Elst (1988), Mann (2000) and from data
collected during this study. Historical data on the numbers
of traps in operation were obtained from aerial photographs
for the period 1949–1978, and thereafter by ground counts
from 1981 to the present.
Results
Fish trapping effort
Information from 1950 to 1994 (Figure 2) shows that the
number of traps and baskets appears to have changed little
during this period, but thereafter there was a substantial
and sustained increase during the late 1990s, particularly
in the number of fish catching baskets, to a peak in 2001.
Subsequently, both trap and basket numbers declined but
remained well above the previous peaks.
Catch size and composition
During the study period a total of 1 759 822 fish was
estimated to have been caught with a mean annual catch
69
Baskets
700 Traps
600
500
400
300
200
100
50
55
60
65
70
75
80
85
90
95
00
05
10
19
19
19
19
19
19
19
19
19
19
20
20
20
YEAR
Figure 2: Numbers of traps and catching baskets in the Kosi Bay
traditional fish trap fishery from 1950 to 2010. Data points prior to
1980 were obtained from aerial photographs and thereafter from
ground counts
of 58 611, though it varied from 28 696 in 1986 to 100 448
in 2006 (Figure 3). The annual number of fish caught in
the traps in the focal study, between 1981 and 1985, was
remarkably constant at around 40 000 and remained close
to this number until 1994 (Figure 3). Subsequently, there
followed a period of erratic but generally much greater
catches until the end of the study.
Although the fish species varied in individual mass, the
165 409 fish caught during the period of intense monitoring
weighed an estimated 161 014 kg, giving a mean fish mass
for this multispecies fishery as 0.97 kg. The mean annual
catch in the early monitoring period was about 40 t.
Fish comprising 43 species from 23 families (Table 2) were
recorded in fish trap catches, but during the study period the
seven most common species were Pomadasys commer-
sonnii, Mugil cephalus, Acanthopagrus vagus, Valamugil
buchanani, Liza macrolepis, Rhabdosargus sarba and Gerres
methueni (Table 2). These combined accounted for almost
95% of the catch by number and 96% by mass (Table 2).
Pomadasys commersonnii and M. cephalus dominated
catches during most of the study period, together contrib-
uting 57.94% by number and 69.71% by mass of the initial
period monitored (Figure 4). Throughout the intensive
monitoring period and in nine of the 11 subsequent years,
P. commersonnii and M. cephalus were the most important
species and this pattern continued up until 1995 when R.
sarba abruptly increased in importance. From a mean of
4.7% of the catch by numbers in the preceding 14 years,
R. sarba rose to 34.7% of the catch in 1995 and, errati-
cally, mostly remained a more important component of
trap catches. Liza macrolepis also changed its contribu-
tion markedly during the study, from being important in the
intensive period to almost disappearing for several years
before returning in numbers in 1999 and becoming the most
important species (by number) at the end of the study.
Although little detailed information is available on
what factors affected annual species catches, Valamugil
buchanani showed a strong negative relationship with
 70 Kyle

All other species/ Rhabdosargus sarba Acanthopagrus vagus Gerres methueni Liza macrolepis Pomadasys commersonnii Mugil cephalus
unknown
105

90
NUMBER OF FISH (×103)

75

60

45

30
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15

1981 1983 1985 1987 1989 1991 1993 1995 1997 1999 2001 2003 2005 2007 2009
YEAR

Figure 3: Yearly catch totals and species composition, by numbers of fish, of the Kosi Bay traditional fish traps from 1981 to 2010. Data for
the period 1981–1984 are total counts and those for 1985–2010 are extrapolations. Note that no data were available for 2001 and 2002

Table 2: Fish catch and species composition, by number and mass, of Kosi Bay fish traps from March 1981 to February 1985 (n = 165 409 fish,
equivalent to 161 014 kg). Field identifications were by Zulu name, other names indicate which species had to be ‘lumped’ within each Zulu name.
The most important five species, in terms of mass, are shown in bold text

Family Species Common name Zulu name Number (%) Mass (%)
Anguillidae Anguilla bicolor Shortfin eel Mbokwane 0 0.01
Anguilla marmorata Mottled eel Ibalane 0.01 0.03
Belonidae Strongylura leiura Garfish Umhlanga 0.11 0.03
Carangidae Caranx ignobilis Giant kingfish Ikomane 1.96 2.36
Caranx papuensis Brassy kingfish Ikomane
Caranx sem Blacktip kingfish Ikomane
Caranx sexfasciatus Big eye kingfish Ikomane
Scomberoides tol Queenfish Umkwandla 0.17 0.13
Chanidae Chanos chanos Milkfish Umnonzi 0.70 1.76
Clariidae Clarias gariepinus Sharptooth barble Ubabule 0.03 0.05
Cichlidae Oreochromis mossambicus Mozambique tilapia Ikwamba 0.92 0.47
Elopidae Elops machnata Springer Ishange 0.33 0.60
Eleotridae Eleotris fusca Dusky sleeper Isimumumu 0.02 0.01
Eleotris melanosoma Broadhead sleeper Isimumumu
Gobiidae Glossogobius giuris Tank goby Isimumumu
Gerreidae Gerres acinaces Smallscale pursemouth Isihlengane 0.35 0.08
Gerres methueni Evenfin pursemouth Inshuludla 9.13 2.57
Haemulidae Pomadasys commersonnii Spotted grunter Inkolonkolo 32.60 36.95
Labridae Thallasoma herbraicum Wrasse Mazinyo 0.03 0.01
Lutjanidae Lutjanus argentimaculatus River snapper Umwali 0.93 2.27
Lutjanus fulviflamma Blackspot snapper Isigalijozi 0.01 0
Monodactylidae Monodactylus argenteus Natal moony Isiphasi 0.04 0
Monodactylus falciformis Cape moony Isiphasi
Mugilidae Crenimugil crenilabris Fringe-lip mullet Isithonga 0.04 0.01
Liza alata Diamond mullet Ibangwa 1.57 2.03
Liza macrolepis Largescale mullet Itefa 10.65 3.41
Mugil cephalus Flathead mullet Intulu 25.19 32.56
Myxus capensis Freshwater mullet Umsiva 0.33 0.11
Valamugil buchanani Bluetail mullet Imbulakazi 2.20 4.27
Valamugil robustus Robust mullet Umacijane 0.10 0.03
Platycephalidae Platycephalus indicus Bartail flathead Umkhokhoba 0.24 0.29
Pomatomidae Pomatomus saltatrix Shad Umuyende 0.19 0.07
Sciaenidae Argyrosomus japonicus Dusky kob Uhlonzi 0.04 0.04
Scorpaenidae Pterios volitans Devil firefish Ugezi 0.01 0
Serranidae Epinephelus andersoni Spotted rockcod Ilondwa 0.07 0.06
Epinephelus gauza Yellowbelly rockcod Ilondwa
Epinephelus malabaricus Malabar rockcod Ilondwa
Sparidae Acanthopagrus vagus River bream Ikula 6.58 4.80
Rhabdosargus holubi Cape stumpnose Isigobolo
Rhabdosargus sarba Natal stumpnose Isigobolo 4.68 3.41
Sphyraenidae Sphyraena barracuda Great barracuda Indobolo 0.74 1.57
Sphyraena jello Pickhandle seapike Indobolo
Theraponidae Therapon jarbua Thornfish Imata 0.02 0
 African Journal of Marine Science 2013, 35(1): 67–78 71

By number By mass 1985 2010

35 30
30

COMPOSITION (%)
COMPOSITION (%)

25
25
20
20
15
15
10
10
5 5

p.
p.
va s

ch a
ac us

et is

R us

ac ta
e

er s
e
er s
V. R. us

Sp . ch ata
ep nn .

L. rae os
ba

L. ae
an tus

A. eni

an ni
ii

.c n .
L. eph ii
.c o P

C cula i
nt eth i
A. halu

O latu
bu arb
M rso P

da
ac da
th pi
a en
ge m an

sp
sp

im la
n

C ana
al

g
g

hy an
bu sar

G role
d

O ole
C al

hu

va

nt . a
gi
m ni
im u
ar G. han

gi

V. . s

u
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M ers

th
e
c

.m

ar
ar
m

m
m

m
co

co

ge
ar
SPECIES SPECIES

L.
L.

Figure 4: Percentage species composition of the more important Figure 5: Catch species composition, by number, of the Kosi Bay
species, by number and mass, of the Kosi Bay traditional fish traps traditional fish traps for 1985 and 2010
from 1981 to 1984

Traps
Table 3: Mean salinity in Makhawulani Lake and yearly trap catches Baskets
of Valamugil buchanani. Correlation coefficient r = 0.988, significant at 1000
p < 0.02
NUMBER OF FISH

800
Mean salinity in Catch of
Year
Makhawulani V. buchanani 600
1981 17.0 517
1982 15.5 687
400
1983 13.4 769
1984 7 1 656
200

salinity measured in Makhawulani Lake during the intensive

study period (Table 3)
90

95

00

05

10
85

19

19

20

20

20
19

With the exception of L. macrolepis, and despite some

YEAR
marked changes, the most important species at the begin-
ning of the study were still very important at the end of it
Figure 6: Estimates of annual catch per fish trap and fish trap
(Figure 5). basket at Kosi Bay from 1981 to 2010

Catch per unit effort

The estimated fish catch per year (Figure 6) showed no clear
overall trend throughout the study, despite the relatively high
peaks, such as in 1995 and 1996, and troughs (e.g. 1986).
No correlations were found between abiotic parameters,
such as annual rainfall, and annual total catch.
Bycatch
Kosi Bay traditional fish traps caught almost exclusively
fish during the intensive monitoring period, with relatively
few catches of mud crabs Scylla serrata (n  76), prawns
(Penaeus spp., n  10), Cape cormorants Phalocrocorax
capensis (n  3), crocodiles Crocodilus niloticus (n 
12) and one green turtle Cholonia mydas. All crocodiles
caught, as well as the green turtle, were translocated
and released away from the fish traps and other human
activity. Cape cormorants are infrequent visitors to the Kosi
Bay lakes (Sinclair 1984) and many of the thousands of
birds that appeared at Kosi Bay in 1994 were caught and
died in the fish traps, as well as in gillnets (Kyle 1996).
Whereas Cowley (1998) reported a mass incursion of Cape
cormorants elsewhere in 1994, which had a serious impact
on fish populations, it appears that at Kosi Bay any such
incursion by Cape cormorants was rapidly depleted through
their capture in fish traps and nets. None of the reed
cormorants Phalacrocorax africanus and whitebreasted
cormorants Phalacrocorax carbo, which are common and
resident at Kosi Bay, were recorded caught in the traps.
Monthly species abundance
All species caught in large numbers in the traps exhibited
monthly patterns in catch abundance. Pomadasys commer-
sonnii and M. cephalus were caught throughout the year,
peaking in mid-summer and late summer respectively
(Figure 7). Catches of G. methueni and A. vagus peaked in
 72 Kyle

(a) P. commersonnii
4 000
3 500
3 000
2 500
2 000
1 500
1 000
500

(b) A. vagus
1 000
800
600
Downloaded by [University of Saskatchewan Library] at 04:06 28 December 2014

400
200

(c) M. cephalus
3 500
3 000
2 500
2 000
1 500
NUMBER OF FISH

1 000
500

(d) G. methueni
1 600
1 400
1 200
1 000
800
600
400
200

(e) Carangidae
300
250
200
150
100
50

(f) Sphyraenidae
140
120
100
80
60
40
20
A M J J A S O N D J F M A M J J A S O N D J F M A M J J A S O N D J F M A M J J A S O N D J F M
1981 1982 1983 1984

Figure 7: Monthly total numbers of (a) Pomadasys commersonnii, (b) Acanthopagrus vagus, (c) Mugil cephalus, (d) Gerres methueni,
(e) Carangidae and (f) Sphyraenidae caught in the Kosi Bay traditional fish traps from 1981 to 1984
 African Journal of Marine Science 2013, 35(1): 67–78 73

mid-summer and in late-summer respectively. Carangidae of which 72% were mature (Kyle 1986). An exception to
and Sphyraenidae showed clear increases in catches this pattern was P. commersonnii, with 30% being <40 cm,
during the summer (Figure 7). the minimum legal size limit. Many fish were also below the
33 cm for male and 39 cm for female length-at-50% sexual
Daily species abundance maturity (Figure 9) (Mann 2000). Almost all Mugilidae
During periods of high catches of individual species, nearly caught were mature and of legal length; however, there is
all showed distinct two-week cycles in abundance. There no size restriction placed on mullet.
was a generally strong relationship between high catches In April 2005, the size and bag limits of many linefish
and maximum daily water height, which were correlated to species were changed. Whereas bag limits should be applied
spring tides measured at Lake Makhawulani. Most species, to fish traps, they have never been as fish that have entered
such as P. commersonnii (Figure 8), were caught in greater the catching baskets can only really be taken out by spearing
numbers in deep water whereas others, such as L. alata, them. The minimum size limits were increased for some
appeared to be caught almost exclusively during dark-moon species, but those of the main species caught in the traps
periods (Figure 8). remained the same. However, by the end of the study, a
much greater proportion of smaller fish were caught for some
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Length frequency analysis species (Figure 9). Each species appeared to show different
During the intensive monitoring period, length frequency
analysis of data from the more important species showed
that few fish <25 cm or >70 cm were caught, and approxi- Mugil cephalus
1981–1984
mately 79% of all fish caught were over the legal size limit, 7
2010
6
(a) 5
Spring tide Spring tide
140 4
120 3
2
100
NUMBER OF FISH (Ƈ) and WATER HEIGHT (cm) (Ƒ)

1
CONTRIBUTION TO CATCH (%)

80
0
60 Water height
20

24

28

32

36

40

44

48

52

56

60

64

68

72

76
P. commersonnii Rhabdosargus sarba
40 10

20 8
0 6
1

10

13

16

19

22

25

28

31

May 1981
4
140 Dark moon Full moon Dark moon Full moon
(b)
2
120
0
100
20

24

28

32

36

40

44

48

52

56

60

64

68

72

76
Pomadasys commersonnii
80 10

60 Water height
8
L. alata catch
40 6

20 4

0 2
25
28
31

3
6
9
12
15
18
21
24
27
30

3
6
9
12
15
18

March April May 0

1984
16
20
24
28
32
36
40
44
48
52
56
60
64
68
72
76

DAYS OF CYCLE TOTAL LENGTH (cm)

Figure 8: Kosi Bay traditional fish trap catch of (a) Pomadasys Figure 9: Length frequency of three important fish species caught
commersonnii for the period 1 May 1981–31 May 1981 and in the Kosi Bay traditional fish traps from 1981 to 1984 and for
(b) Liza alata for the period 25 March–19 May 1984 and water 2010. The length at sexual maturity is indicated (downward arrow)
height measured at Lake Makhawulani. Spring tides were on 4 and as well as the legal minimum length for capture (upward arrow).
18 May 1981 and 1, 15 and 29 April and 13 May 1984; new moons Mugil cephalus does not have a legal minimum length
were on 4 May 1981 and 1 and 29 April 1984
 74
degrees of change in length frequency between the early and
later stages of the study. Some fish, such as M. cephalus,
showed very similar patterns in the earlier and later phases
of the study whereas others showed a small decline (R.
sarba) or marked decline (P. commersonnii) in mean length.
Mugil cephalus has recently become an important
species in fisheries and aquaculture worldwide (Whitfield et
al. 2012), but with wild catches reportedly declining in many
areas in South Africa. Due to its apparently fairly stable
mean size in trap catches (Figure 9) and reported ability to
survive without using estuaries (Whitfield et al. 2012), M.
cephalus would be more likely to withstand heavy fishing
pressure than other fish. Thus, the apparent decline in their
proportion in the trap catches, from 25.6% during the first
five years of the study to 18.2% in the last five years, is
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cause for concern.
Discussion
The most complete data presented here are those obtained
from the 48 months (March 1981–February 1985) when all
fish traps were monitored. Extrapolations of total catch from
a single area provide a useful insight into overall total catch
numbers but species composition will be skewed in favour of
the more abundantly caught species in the areas monitored.
Information from the early monitoring period suggested that,
for example, traps near the mouth sometimes had a different
catch composition than that from traps in Lake Makhawulani.
Kosi Bay fish traps are set in shallow water, where there
are few large fish during the day. Fish are caught mainly
at night, and the fact that many of the fish have well
developed or developing gonads (Kyle 1986) suggests that
many fish caught could be on spawning-related migrations.
Kyle (1986) found a highly significant correlation between
gonad development and increased catches in species
such as M. cephalus and A. vagus. The author found that
A. vagus caught during peak catches had well-developed
gonads. Spawning of most of the important trap-caught
species takes place at sea (van der Elst and Adkin 1991)
or just inside the Kosi mouth (Garratt 1992). It is likely that
most fish caught in the traps are moving seaward as most
traps face upstream. Most traps were built facing upstream
because the trappers believe that virtually only seaward-
moving fish could or should be caught.
Catch size and species composition
In terms of overall catch, the traps caught similar numbers
and mass of fish annually over the intensive study period.
Historical data suggest that trap numbers have also been
fairly constant in living memory and up until about 1994
(Figure 2). It thus seems likely that total trap catches,
historically and up until recently, represented a fairly stable
proportion of the fish population of the lake’s system
each year. Data from a tagging experiment (Kyle 1986)
suggested that the traps caught between 5% and 10% of
the total adult population of most large fish species annually
(Figure 10). Subsequent mark and recapture supports this
estimate and it appears that, until about 1990, the traps
were a fairly stable and easily managed method to capture
fish. The overall increase in trap numbers, and particularly
the baskets (Figure 2), as well as the estimated overall
Kyle
35
30
RECOVERED (%)
25
20
15
10
5
83
19 85
08
9
4
05
06
07
09
10
–9
–0
19
19
20
20
20
20
20
20
89
00
YEAR/ PROJECT / MEAN
Figure 10: Percentage of marked fish recovered at Kosi Bay by
traditional fish traps and other fishing methods from 1983 to 2010.
Data for 1983 and 1985 were from specific projects, 1989–1999
are the mean annual recovery rates and 2004–2010 are the annual
marked fish recovery rates each year
marked increase in catches, need to be monitored in case it
leads to greatly increased and consequently unsustainable
catches. Kyle (1986), however, suggested that providing a
trap-free 30 m channel was kept open between the lakes
and the sea, the proportion of fish ‘straying’ into the traps
would be similar regardless of the number of traps. All
trap sites were different and those that consistently had
good catches were consistently occupied and maintained
in good condition over many years. Less productive traps
were often abandoned and they slowly disintegrated and
eventually were washed away. When the number of traps
increased markedly, they often had to be built in progres-
sively less productive sites. Continued mark and recovery
results (Figure 10), however, appear to show a progressive
increase in the proportion of tagged fish being recovered.
While it seems likely that earlier recovery rates were indeed
initially sustainable, the later rates suggest that this situation
may have changed.
Total trap catches subsequent to the intensive study were
more variable than those between 1981 and 1985, which is
to be expected when only a proportion of the fishing area
is monitored and extrapolations had to be made. Changes
in the numbers of some species caught in the later stage
of the study (e.g. A. vagus) are probably due to different
species being more abundant in different areas, depending
on salinity and other abiotic factors. Salinity is an important
factor in the distribution of some fish species (Blaber 1973,
Whitfield et al. 1981, Cyrus and Blaber 1982, Whitfield
1998) and the response of closely related species to similar
salinity regimes can vary markedly (Cyrus and Blaber
1982).
It is likely that the total annual trap catch numbers result
from such diverse factors as spawning success, weather
conditions, and natural and man-induced mortality levels.
While some of these factors act near or within the Kosi Bay
system, others may be more general oceanic or climatic
effects.
 African Journal of Marine Science 2013, 35(1): 67–78
Access to, and recruitment from, other estuarine systems
could be an important factor in controlling the numbers of
some fish species. The St Lucia Lake system, the largest
estuarine system in southern Africa, is less than 200 km
away from the Kosi Bay system, but it has effectively been
closed to the sea for over 10 years. Species such as R. sarba
are reported to spawn in the nearshore environment (Mann
2000), but juveniles generally enter estuaries where they
grow to maturity before moving to the open ocean. During
the past decade, catches of this species have been high in
traps and it is probable that the closure of St Lucia resulted in
an increase of recruits entering the Kosi Bay system as they
would be unable to enter the St Lucia Estuary.
Overall, there was a general increase in trap catches over
the study period that seems to correspond to the increase
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in trap numbers. The CPUE was variable and no pattern
was discernible during the study. This could be related to
changes in the traps themselves and the materials used.
An underlying ‘driver’ in the trap fishery is that it used to
be at subsistence level, but positive developments of a local
economy and better access to markets have changed this.
In the past, if a trapper caught a lot of fish he often shared
them with neighbours and friends. In the current situation,
if a lot of fish are caught in the traps, the owner often sells
the entire catch to a person who exports the fish from the
region. This practice not only takes relatively cheap protein
away from much of the local community but it opens a
consistent lucrative market for trap fish. A 3 kg P. commer-
sonnii was sold in August 2012 in KwaNgwanase market for
R180, and in 2010 the same species of similar size would
have been R90 (i.e. R30 kg–1). An estimate of the value
of the 2010 catch would thus be R 2 250 000 and, as all
sizes of fish are sold at similar prices per kg, there is every
incentive for trappers to catch as many fish and of any size
as possible.
Seasonality
All species caught in large numbers in the traps showed
annual cycles of abundance, many of which appeared to
be associated with their breeding cycles. For example,
the numbers of A. vagus peak in the months preceding
their spawning, just inside the mouth of the Kosi (Garratt
1992). Other catches appeared to reflect the immigration
and emigration of fish that are reported to enter estuaries
in early summer and exit in late summer (Wallace 1975).
These fish are mainly Carangidae, Sphyraenidae and L.
argentimaculatus, which feed mainly on fish, but some are
detritivores (e.g C. chanos).
Fish traps are operated throughout the year but catches
of most species peak in the warmer summer months.
There is usually a clear increase in catches of G. methueni
in mid-summer followed by a slightly later peak in A.
vagus catches. Although caught throughout the year, M.
cephalus tends to be in greater numbers in late summer.
Pomadasys commersonnii catches are highest in middle–
late summer, often associated with heavy rains that would
result in increased water flow and also darker water due
to tannins being washed into the lakes (RK pers. obs.).
These latter two species are the most important to the
trappers and trap maintenance is often manipulated to
catch them during their predictable annual ‘runs’. Catches
75
of some species fluctuated markedly from one year to
the next, and appeared unrelated to any obvious factors
(Figure 3), whereas other species showed a high degree
of stability in the catch rates. The species that showed
large fluctuations in annual catch were those whose life
cycles were spent mostly outside estuaries, such as
Carangidae and Sphyraenidae. More estuarine-associated
species, such as A. vagus, whose catches varied only up
to a maximum of 20% around the mean per year during
the intensive study, as well as G. methueni, exhibited a
higher degree of catch stability. Few A. vagus have been
recorded in the open sea off the East African coast and
Garratt (1992) recorded them spawning inside the Kosi
mouth. Several hundred A. vagus were tagged inside the
lakes system (Kyle 1986), but no recoveries were made
from the sea, which suggests that this species is restricted
mainly to the Kosi Bay lake system.
Of the approximately 5 000 fish tagged inside the Kosi
mouth during the entire study, only one fish was recovered
at sea and it was released and recovered again back inside
Kosi Bay. Only one fish (an M. cephalus) recovered from
a fish trap was tagged outside Kosi Bay, in an estuary in
the Western Cape. Overexploitation by the traps or other
methods within the lakes system of heavily estuarine-
related species at Kosi Bay will probably result in future trap
and other catches of those species. Future catches of other,
less estuarine-dependent species, including P. commer-
sonnii and M. cephalus, may be more independent of fish
trapping pressure as it appears that recruitment comes from
a much larger population outside the Kosi Bay lakes.
Over the period 1981–1984, there was a high correla-
tion (r  0.995) between P. commersonnii trap catches and
catch rates from shore-fishing competitions along the KZN
coast (Kyle 1986). This suggests that the same parameter
modified the catches of this species along much of the KZN
coast.
Daily fluctuations
Within the annual periods of catch abundance there were
usually distinct 14-day cycles (Figure 8). These were associ-
ated with spring tides and the resultant high water levels, but
this pattern may be distorted or modified by heavy winds or
rain that can affect water level. During periods of increased
catches of a particular species, there was a strong correla-
tion between water depth at Lake Makhawulani and total
trap catch on a daily basis (Figure 9, Kyle 1986). On 28
February 1984, cyclone Domoina provided insight into the
impact of particularly high water levels as the best catches
were recorded for several species, including P. commer-
sonnii, on the days following this event when water levels
were particularly high. Kyle (1986) found the strongest
relationship between trap catches and daily water depth
when considering various biotic and abiotic variables. At
Kosi Bay, the turbidity in the northern channels is usually
low, even during moderate floods, but large-scale flooding
usually washes dark tannins into the channels, which also
appears to promote the movement of some fish species.
Conservation and management
The monitoring programme presented here was imple-
mented because of concern of the possible impact of trap
 76
catches on fish populations and thus on the catches of
other fishing sectors, particularly the recreational anglers.
In the early stages of the study, up to about 1994, it
appeared the annual trap catch was a fairly stable propor-
tion of fish moving to the sea, and catches were low
enough to allow adequate fish to exit the lakes system
or reach the spawning grounds. Kyle (1986) showed that
during the intensive study period, 52% of the total trap
catch by number and 47% by mass were of mullet G.
methueni and other species not important to recreational
or other rod-and-line anglers. The fairly low proportion of
the fish population caught in the traps, estimated at about
5% of fish annually (Kyle and Robertson 1997), and the
species composition suggested that, at that time, trap
catches were sustainable and not a serious threat to the
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recreational and other fishing sectors.
Subsequent to 1994, however, there was a sustained
increase in trap catches coupled with a marked and
progressive increase in the proportion of marked fish being
recaptured. Bearing in mind that marked fish recovery
information produces a minimum estimate of impact on
stocks due to emigration and mortality of marked fish,
tag-loss, underreporting of marked fish caught and other
aspects, a reported recovery rate of about 35% (Figure 10)
may well reflect a very high and unsustainable impact on
fish stocks.
A further conservation concern is that the traps were
catching many fish that were immature or less than the legal
minimum length. Several of the most important species,
such as M. cephalus, G. methueni and L. macrolepis, are
not ‘linefish’ and thus do not have ‘legal’ size limits. Analysis
of the data from the intensive study period and the final year
of the study of some important species (Figure 9) show that
while all the M. cephalus were of legal length, and mostly
of mature size, large numbers of small and often immature
R. sarba and P. commersonnii were being caught. Although
the proportion of small R. sarba caught remained similar
between the beginning and end of the study, by 2010
total catches of this species were much greater. Also, the
capture of an increasing number of undersize and immature
P. commersonnii must be of conservation management
concern. A possible explanation for the marked decrease in
the length of P. commersonnii between the earlier and later
data could be on account of the recent practice of facing
some traps downstream rather than upstream, which is the
traditional method of fishing. Traps facing downstream would
catch the small recruits entering the system as opposed to
larger fish exiting the system. If this is shown to be the case,
then such trapping methods should not be used in the future.
With Lake St Lucia being effectively closed to the sea for
many years, and thus having a negligible ‘nursery function’
over this period, this renders Kosi Bay more important in
terms of providing recruits to fish populations. Species such
as R. sarba and P. commersonnii are important in the KZN
recreational shore-fishery and so marked reductions in fish
exiting the Kosi Bay lakes will impact this important fishery.
Most of the Carangidae, Sphyraenidae and Lutjanidae
were immature but of legal size (van der Elst 1988). Using
data from Figure 9 and from Kyle (1986), it is estimated
that during the intensive study period, 84.6% of the total
catch by numbers were legal in terms of the current linefish
Kyle
legislation, and 74.2% of the fish for which adequate length-
at-maturity data are available were greater than the length-
at-50% maturity (van der Elst and Adkin 1991). By the end
of the study, however, this situation had changed markedly,
an estimated >70% of P. commersonnii being immature and
an even greater proportion below the legal size limit.
Skelton (1987) identified at least eight threatened fish
species present in the Kosi Bay lakes. Most of these are
too small to be caught in traps; however, freshwater mullet
Myxus capensis is caught frequently and the broadhead
sleeper Eleotris melanosoma is caught in appreciable
numbers. The capture of an estimated 1 885 M. capensis
in 1995 within a protected area might be seen as cause for
concern, but if it reflects a stable and sustainable proportion
of species stocks annually, then no action needs to be taken
to protect this species. If, however, as suggested by the
most recent data from this study (Figure 10), trap catches
have increased and traps are now catching unsustainable
numbers of estuarine-dependent species, then management
action would be necessary, at least for these fish.
Kyle (1986) suggested that, as long as the 30 m trap-free
channel was maintained, the number of traps was not
directly related to the total catch, because the best trap
areas were those first occupied and newer traps were in
less productive areas. Our study showed no clear trend
in trap CPUE over the 30 years of monitoring, despite
an overall increase in trap numbers. This pattern of an
increase in the number of traps leading to larger total
catches indicates that a continued increase in trap numbers
would be unsustainable. In the past few years, however,
fish traps have sometimes been placed in the trap-free
channel (RK pers. obs.) and modern materials have been
used in their construction. This resulted in these traps
becoming more efficient at catching fish and thus increasing
overall trap catches. This in turn would reduce the value of
CPUE as an indicator of the health of the fish stock.
Whereas building traps into the channels would increase
the proportion of fish caught, the use of modern materials
would enable the traps to catch smaller fish. Many trappers
now use nylon rope and gumpoles to reduce the gaps
between the sticks in the terminal baskets in order to
retain smaller, and often immature, fish. Historically, fish
trapping was a low-key method of supplementing income
and providing protein for the trappers’ community. Recently,
however, it has become increasingly commercialised with
the result that trap catches are now driven by economic
incentive and some of the traditional constraints have been
abandoned. As fish are generally sold per kg, even small
fish such as the thornfish Terapon jarbua are now routinely
sold in local markets.
Traps are efficient as there is only a small bycatch.
Hippopotomus still occur in appreciable numbers in the
lakes and are not negatively impacted by the traps although
occasionally they damage traps by walking through them.
Recreational anglers catch fish in the lakes before they enter
the trap areas and so their catch is not affected by the traps.
Also, many of the species caught in traps are not targeted by
recreational anglers. This means that traps, except for those
that face downstream, will have minimal short-term impact
on recreational catches. Kyle (1986) estimated that recrea-
tional anglers caught about 1% of adult fish stocks annually
 African Journal of Marine Science 2013, 35(1): 67–78
and their catches appear to have remained fairly stable
throughout the past three decades and so their catches
should not impact markedly on trap catches.
From this study, it appears that the initial perceived conflict
between traditional fish trappers and recreational anglers
is unsubstantiated. Neither of the two fisheries appears
to be seriously impacting on the other and both appear to
be harvesting fish at sustainable levels. More recently,
however, the rising numbers of fish being caught (especially
immature fish of important species) and the increasing
efficiency of the traps have raised their offtake proportion
markedly to levels that are likely to be unsustainable.
While traditional trapping and other fisheries in Kosi
Bay likely coexisted sustainably in the past, there is now
compelling evidence that overall catch rates for many
Downloaded by [University of Saskatchewan Library] at 04:06 28 December 2014
species are unsustainable. Management intervention is now
required to reduce overall fishing effort and to monitor the
changes in the the construction and thus efficiency of the
traps. Action is now required to reduce catches to levels
that appear to be wise, sustainable and appropriate for this
World Heritage Site, to the mutual benefit of local trappers
and other fishers.
Acknowledgements — I thank my family for help and support
during this study and Ezemvelo KZN Wildlife for permission to
publish this paper. Thanks are due to the people who participated
in the data collection and for maintaining good relations with the
fish trappers.
References
Begg G. 1978. The estuaries of Natal. Natal Town and Regional
Planning Report 41.
Begg G. 1980. The Kosi system: aspects of its biology, manage-
ment and research. In: Bruton MN, Cooper KH (eds), Studies on
the ecology of Maputaland. Grahamstown: Rhodes University. pp
358–373.
Beliaef B, Louis M, Souprayen J. 1992. Preliminary results on the
efficiency and selectivity of the chevron-shaped trap, a traditional
fishing device in Guadeloupe. Proceedings of the Gulf and
Caribbean Fisheries Institute 41: 150–165.
Blaber SJM. 1973. Temperature and salinity tolerance of juvenile
Rhabdosargus holubi (Teleostei: Sparidae). Journal of Fisheries
Biology 5: 593–599.
Blaber SJM. 1978. Fishes of the Kosi system. Lammergeyer 24:
28–41.
Blaber SJM. 1981. A revised checklist and further notes on the
fishes of the Kosi system. Lammergeyer 31: 5–14.
Copes P. 2000. Aboriginal fishing rights and salmon management
in British Columbia: matching historical justice with the public
interest. In: Knudson EE, Steward CR, MacDonald DD, Williams
JE, Reiser DW (eds), Sustainable fisheries management: Pacific
salmon. Boca Raton, Florida: CRC Press LLC. pp 75–94.
Cowley PD. 1998. Ichthyofaunal population dynamics of the
East Kleinemonde Estuary, South Africa. PhD thesis, Rhodes
University, South Africa.
Cyrus DP. 1980. The biology of the Gerreidae Bleeker, 1859
(Teleostei) in Natal estuaries. MSc thesis, University of Natal,
Pietermaritzburg, South Africa.
Cyrus DP, Blaber SJM. 1982. Species identification distribution
and abundance of Gerreidae (Teleostei) Bleeker, 1859, in the
estuaries of Natal. South African Journal of Zoology 17: 105–116.
Cyrus DP, Blaber SJM. 1984a. Predation and sources of mortality
of Gerreidae Bleeker, 1859 (Teleostei), in Natal estuaries with
special reference to the Kosi system. Lammergeyer 32: 14–20.
77
Cyrus DP, Blaber SJM. 1984b. The feeding ecology of Gerreidae
Bleeker, 1859 (Teleostei) in the Kosi system, with special
reference to their seasonal diet. Lammergeyer 32: 35–49.
Felgate WS. 1965. An ecological study of the Tembe Thonga of
Natal and Mozambique. Unpublished report on research work
edited by EJ Krige. University of the Witwatersrand, South Africa.
Fennessy ST, Everett B, Radebe V. 2001. Stock assessment and
management of the spotted grunter in KwaZulu-Natal (final
report). Unpublished Report No. 200. Durban: Oceanographic
Research Institute.
Garratt PA. 1992. Spawning of riverbream, Acanthopagrus berda,
in Kosi estuary. South African Journal of Zoology 28: 26–31.
Hill BJ. 1969. The bathymetry and possible origins of Lakes Sibaya,
Nhlange and Sifungwe in Zululand (Natal). Transactions of the
Royal Society of South Africa 38: 205–216.
James NC, Beckley LE, Mann BQ, Kyle R. 2001. The recreational
fishery in the Kosi estuarine lake system, South Africa. African
Zoology 36: 217–228.
Koranteng KA, Ofori-Danson PK, Entsua-Mensah M. 2000.
Fish and fisheries of the Muni lagoon in Ghana, West Africa.
Biodiversity and Conservation 9: 487–499.
Kyle R. 1981a. The fish kraals of the Kosi system. African Wildlife
36: 186–187.
Kyle R. 1981b. The golden sleeper fish – extension of range.
Lammergeyer 31: 43.
Kyle R. 1986. Aspects of the ecology and exploitation of the fishes
of the Kosi Bay lakes system, KwaZulu-Natal, South Africa. PhD
thesis, University of Natal, Pietermaritzburg, South Africa.
Kyle R. 1996. A mass incursion and mortality of the Cape cormorant
Phalocrocorax capensis, in the Kosi Bay lakes. Lammergeyer 44:
43–45.
Kyle R, Robertson W. 1997. Preliminary estimates of population
size and capture rates of mature Acanthopagrus berda in the
Kosi lakes system, South Africa, using mark-recapture methods.
South African Journal of Science 32: 124–128.
Mahon R, Drayton N. 1990. Trap fishery management in Barbados:
the fisherman’s perspective. Caribbean Marine Studies 1: 94–100.
Mann BQ. 2000. Southern African marine linefish status reports.
Special Publication No. 7. Durban: Oceanographic Research
Institute.
Mountain A. 1990. Paradise under pressure. Cape Town: Southern
Book Publishers.
Mwatha GK, Orembo B. 1998. Fishery in Mida Creek. In: Mwatha,
GK, Fondo E, Uku JN, Kitheka JU (eds), Biodiversity of Mida
Creek: final technical report. Mombasa: Kenya Marine and
Fisheries Research Institute. pp 114–134.
Russ GR, Acala AC. 1996. Do marine reserves export adult fish
biomass? Evidence from Apo Island, Central Philippines. Marine
Ecology Progress Series 132: 1–9.
Shaul W, Reifsteck D. 1991. Comparison of catches by bamboo
and wire fish traps in Jamaica. Proceedings of the Gulf and
Caribbean Fisheries Institute 40: 99–107.
Siddeek MSM, Fouda MM, Hermosa GV. 1999. Demersal fisheries
of the Arabian Sea, the Gulf of Oman and the Arabian Gulf.
Estuarine, Coastal and Shelf Science 49: 87–97.
Sinclair JC. 1984. Field guide to the birds of southern Africa. Cape
Town: Struik.
Skelton PH. 1987. South African red data book – fishes. South
African National Scientific Programmes Report 137. Pretoria:
National Scientific Programmes Unit: CSIR.
Tinley KL. 1964. Fishing methods of the Thonga Tribe in north-eastern
Zululand and southern Mozambique. Lammergeyer 3: 9–39.
van der Elst RP. 1988. A guide to the common sea fishes of
southern Africa. Cape Town: Struik.
van der Elst RP, Adkin F. 1991 Marine linefish priority species and
research objectives in southern Africa. Special Publication No. 1.
Durban: Oceanographic Research Institute.
 78
Wallace JH. 1975. The estuarine fishes of the east coast of South
Africa. South African Association for Marine Biological Research.
Investigational Report 40.
Whitfield AK. 1998. Biology and ecology of fishes in southern
African estuaries. Ichthyological Monographs of the JLB Smith
Institute of Ichthyology 2.
Whitfield AK, Blaber SJM, Cyrus DP. 1981. Salinity ranges of
Downloaded by [University of Saskatchewan Library] at 04:06 28 December 2014
Kyle
some Southern African fish species occurring in estuaries. South
African Journal of Zoology 16: 151–155.
Whitfield AK, Panfili J, Durand JD. 2012. A global review of the
cosmopolitan flathead mullet Mugil cephalus Linnaeus 1758
(Teleostei: Mugilidae), with emphasis on the biology, genetics,
ecology and fisheries aspects of this apparent species complex.
Reviews in Fish Biology and Fisheries 22: 641–681.
Received June 2012; accepted October 2012