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Estuarine, Coastal and Shelf Science (2000) 50, 259–269

doi:10.1006/ecss.1999.0560, available online at http://www.idealibrary.com on

Aspects of the Biology of Four Species of Sciaenidae


from the East Coast of South Africa
S. T. Fennessy
Oceanographic Research Institute, P.O. Box 10712, Marine Parade, 4056, South Africa

Received 29 March 1999 and accepted in revised form 25 August 1999

The biology of four Indo-Pacific sciaenid species, Otolithes ruber (Schneider, 1801), Atrobucca nibe (Jordan & Thompson
1911), Johnius dussumieri (Cuvier, 1830) and Johnius amblycephalus (Bleeker, 1855), occurring off the KwaZulu-Natal
coast, is described. Samples were collected from prawn trawlers, beach-seine netters and boat anglers from May 1989 to
February 1994. Biological data include: length composition, length-weight relationships, sex ratios, spawning seasons,
sizes at 50%-maturity and feeding. All four species are relatively small, particularly those individuals caught by
shallow-water prawn trawlers. Females outnumbered males in all four species. Spawning seasons are lengthy, occurring
in spring and summer months. Most of the fish caught by shallow-water trawlers are below their size at 50%-maturity.
The results show that shallow, turbid, continental shelf waters are nursery areas for these four sciaenids.
 2000 Academic Press

Keywords: biology; Sciaenidae; nursery; prawn trawling; bycatch; South Africa; western Indian Ocean

Introduction et al., 1994), biological data were collected for four of


the above species, i.e. O. ruber, A. nibe, J. dussumieri
Representatives of the family Sciaenidae (croakers or and J. amblycephalus. These data are analysed and
drums) are widely distributed in shelf waters of the presented here as a description of aspects of the life
tropical and subtropical Indian, Pacific and Atlantic histories of these fishes in South African east coast
oceans (Lowe-McConnell, 1962; Druzhinin, 1974; waters. The data will also complement recently-
Trewavas, 1977; Longhurst & Pauly, 1987; Sasaki, completed studies on other South African sciaenids
1996) and are important components of fisheries in (Griffiths, 1996a; Griffiths, 1996b; Griffiths, 1997).
several countries (Fischer & Bianchi, 1984). There are
at least 10 sciaenid species occurring in South African
Methods
waters, three of which are important in the hook and
line fishery in the east coast province of KwaZulu-
General
Natal (Mann-Lang et al., 1997). These are Argyro-
somus japonicus, Atractoscion aequidens and Argyrosomus Samples of the four species were collected at irregular
thorpei. Five other sciaenids are also caught by various intervals and on an opportunistic basis from prawn
fisheries in KwaZulu-Natal. Otolithes ruber is targetted trawlers, beach-seine netters and boat anglers on the
by boat anglers off Durban (Beckley & Fennessy, KwaZulu-Natal coast from May 1989 to February
1996) and is a major component of shallow-water 1994. Most samples were from shallow-water
(<50 m) prawn trawl bycatch (Fennessy et al., 1994). (<50 m) prawn trawlers operating on the Tugela Bank
Atrobucca nibe is caught as a bycatch by deep-water trawl grounds approximately 100 km north of Durban
(>350 m) prawn trawlers off Durban (Fennessy & (Figure 1). Atrobucca nibe were also obtained from
Groeneveld, 1997), and Johnius dussumieri and J. deep-water (>300 m) prawn trawlers off Durban
amblycephalus are components of shallow-water prawn (Figure 1). Once the prawns had been removed from
trawler bycatch (Fennessy et al., 1994). The former the trawl samples, a random subsample of between 10
species is also caught by anglers in KwaZulu-Natal and 20 kg of the bycatch was collected with a shovel.
estuaries (Mann, 1993). Finally, Umbrina ronchus is Specimens of the four species present in this sub-
targetted by KwaZulu-Natal spearfishers, albeit in sample were treated according to the procedures
small quantities (Mann et al., 1997). described below. An additional random subsample of
During an investigation of the ichthyofaunal 20 specimens of each species per trawl was frozen on
bycatch of KwaZulu-Natal prawn trawlers (Fennessy board in order to collect information on individual

0272–7714/00/020259+11 $35.00/0  2000 Academic Press


260 S. T. Fennessy

St Lucia Estuary

Tu
ge
la

m
Ri 29°00' S

50
ve
r

0m
20
Tugela Bank

Kwazulu-Natal

Durban
Indian Ocean

30°00' S

South Africa

31°00' S

Trawling grounds

31°00' E 32°00' E
F 1. Map of the KwaZulu-Natal coast showing areas mentioned in the text.

weights and gonad weights in the laboratory. As it Additional data were obtained for J. amblycephalus
became apparent during the first year of data collec- from beach-seine netters in Durban during 1992
tion that numbers of mature fish obtained by random and for O. ruber from catches made by small boat
sampling were low, these samples were supplemented anglers in the same area in 1993 and 1994. The
by the selection of larger fish from the trawl catches. data from fish selected from trawls, and from fish
Biology of Sciaenidae from South Africa 261

T 1. Macroscopic gonad maturity stages utilised to stage sciaenids from KwaZulu-Natal

Stage Description

Inactive Gonads are initially transparent, thread-like and less than half the body cavity in
length. Later in this stage, the gonads are thicker; ovaries are pale yellow-orange,
with no eggs visible and testes are off-white, with no milt in the duct.
Active Gonads are about three-quarters of the body cavity in length. Ovaries are slightly
swollen, darker yellow-orange, with eggs initially just visible, becoming more so later
in the stage. Testes are firm, white and triangular in cross-section; milt occurs in the
spermatic duct.
Ripe Ripe ovaries are very swollen, containing large numbers of eggs which, in spawning
fish, are hydrated and flow freely from the vent. Ripe testes have milt flowing at the
slightest pressure and tend to rupture easily.
Spent Ovaries are much reduced in volume, are reddish and contain some opaque eggs.
Testes are mottled, brownish and much reduced in volume.

collected from beach-seine and boat anglers, were xmid =the midpoint of the class interval;
only used for the calculation of length-weight x0·5 =size at 50%-maturity;
equations, gonado-somatic indices and sizes at 50%- d=the width of the maturity ogive (Butterworth et al.,
maturity. 1989).
Precision of the estimates is given as 95% confidence
intervals based on asymptotic standard errors.
Population composition
Total lengths (mm) of the first 50 undamaged fish of
each of the four species were recorded from each Feeding
sampled trawl. At least 20 fish of each species from Stomachs from at least 20 fish per species per trawl
each sample were sexed to calculate sex ratios. Indi- sample were examined. Stomach contents were ident-
vidual weights (g) were mostly collected during ified to species level wherever possible. Assessment
the first year of sampling in order to calculate was based on the frequency of occurrence method
length-weight relationships. (Ricker, 1971), in which the number of stomachs in
which a food category appeared was recorded.
Gonad development
The maturity of at least 20 fish of each species from Results
each sample was assessed using a macroscopic staging
index (Table 1). Seasonality of spawning was deter-
General
mined by plotting the monthly frequency of each
gonad stage per size class in females and by calculating A total of 110 trawl samples was collected from the
gonado-somatic indices (GSI) for female fish larger Tugela Bank from May 1989 to June 1992. Sampled
than the size at maturity. Indices and numbers per depths ranged from 20 to 50 m, and samples were
gonad stage were pooled within calendar months obtained in all calendar months except December.
among years. The four species investigated (O. ruber, A. nibe, J.
Dussumieri and J. amblycephalus) were not always all
GSI=(weight of gonad/weight of fish)100 represented in catches. An additional 10 trawl samples
of A. nibe (n=975) were collected from deep-water
Size at 50%-maturity estimates were based on female
prawn trawlers from November 1992 to December
fish collected during spawning season, and estimated
1993. Codend mesh size was 38 mm (stretched) in all
from fitted logistic curves of the form:
sampled trawl catches. Eight samples of O. ruber
y=1/(1+exp((xmid x0·5))/d) (n=82) were obtained from Durban boat anglers from
September 1993 to February 1994. Three samples of
where J. dussumieri (n=3) and J. amblycephalus (n=21) were
y=the proportion of mature fish in length class x obtained from Durban beach-seine netters during
during spawning season; December 1992.
262 S. T. Fennessy

(a) positively skewed. There is a significant increase in the


40 mean sizes of all four species with increasing depth on
Trawling n = 3378
the Tugela Bank (Table 2). Overall, females signifi-
30 Angling n = 82
cantly outnumbered males in all four species caught
by trawling (Table 3), particularly in the larger size
%

20
classes (Figure 3). The a and b parameters of the
10
length-weight equations (Table 4) are very similar for
0 O. ruber and A. nibe and for J. amblycephalus and
J. dussumieri, respectively, a reflection of the similar
15

65

5
11

16

21

26

31

36

41

46

51
TL (mm)
morphometrics of the two species pairs.
(b)
40 Gonad development
30 Shallow n = 2073
Deep n = 975 The gonads of all four species were readily assigned
maturity stages according to Table 1. Mature female
%

20
fish did not always occur in monthly catches, which
10 resulted in gaps in the monthly data. Peaks in the
gonado-somatic indices of females (Figure 4) and the
0
increased frequency of ripe fish (Figure 5) indicated
15

65

5
11

16

21

26

31

36

41

46

51

an increase in gonadal activity and protracted spawn-


TL (mm) ing in spring and summer (September to February)
(c)
14 for O. ruber, J. amblycephalus and J. dussumieri. In A.
12 n = 3838 nibe, spawning also occurred in August and March.
10 Observation of partly-spawned and spent female fish
8 in some of these months (Figure 5), assisted in the
%

6 confirmation of seasonality of reproduction.


4 Plots and estimates of the female lengths at 50%-
2 maturity are presented in Figure 6 and Table 5
0 respectively. The high R2 values indicate a relatively
35

55

75

95

good fit to the data for all four species.


11

13

15

17

19

21

TL (mm)
(d)
20 Feeding
n = 3429
15 A total of 1713 O. ruber stomachs, 685 stomachs of
A. nibe, 1319 stomachs of J. dussumieri and 1244
%

10 stomachs of J. amblycephalus were examined. In all


four species, 60 to 70% of stomachs were either empty
5
or exhibited barotrauma (Table 6). Crustacean prey
0 were dominated by brachyuran crabs, caridean
shrimps and penaeid prawns, particularly the small
35

55

75

95

5
11

13

15

17

19

21

23

TL (mm) penaeid, Parapenaeopsis acclivirostris. The Natantia


F 2. Length frequency distributions of four sciaenid (prawns and shrimps) were particularly important in
species from KwaZulu-Natal (May 1989 to February 1994). stomach contents of three of the fish species exam-
Dotted lines indicate female sizes at 50%-maturity. (a) O. ined. Although teleosts were more important in
ruber; (b) A. nibe; (c) J. dussumieri; (d) J. amblycephalus. A. nibe, 60% of stomachs which contained prey were
from larger fish caught by deep-water trawlers. This
Population composition may account for the reduced importance of crusta-
Length frequency distributions indicate that the four ceans in this species. Teleosts and brachyurans were
species are all relatively small (Figure 2). Relative to also important in the diet of O. ruber.
the Tugela Bank trawlers, the boat anglers and the
deep-water prawn trawlers caught larger individuals of
Discussion
O. ruber and A. nibe, respectively. With the exception
of J. dussumieri, the distributions of the sciaenids Publications on the biology of South African sciaenids
in Tugela Bank trawl catches are unimodal and are mostly limited to the genera Atractoscion and
Biology of Sciaenidae from South Africa 263

T 2. Single-factor ANOVA of lengths of four species of sciaenids caught at three depth ranges by Tugela Bank prawn
trawlers (May 1989 to February 1994)

20–33 m 28–33 m 33–45 m


Mean STD n Mean STD n Mean STD n F

O. ruber 149·8 52·9 1513 194·3 67 439 262·3 80·1 679 742·2*
A. nibe 112·1 22·6 373 150·5 65·1 378 176 70·6 500 127·6*
J. dussumieri 114·9 26·9 734 125·5 31·5 841 131·2 31·5 666 63·7*
J. amblycephalus 119·1 35·2 818 126·5 44·2 723 144·5 43 835 84·1*

Fcritical =3 and P<0·001 in each case. *Denotes significance at the 95% level.

T 3. Overall sex ratios of four species of Sciaenidae trawled on the KwaZulu-Natal coast (May
1989 to February 1994)

Species Females Males Ratio 2 P

O. ruber 972 534 1:1·82 127·4 0·001


A. nibe 716 202 1:3·55 287·8 0·001
J. dussumieri 758 604 1:1·25 17·4 0·001
J. amblycephalus 665 589 1:1·13 4·6 0·05

Degrees of freedom (df)=1 in each case.

Argyrosomus (e.g. Wallace & Schleyer, 1979; Smale, 1972), but visual appraisal of the gonads did not
1985; Beckley, 1990; Griffiths & Hecht, 1995a; provide evidence for this. Size disparity between sexes
Griffiths & Hecht, 1995b), largely because of their could also be explained by differential growth and
commercial importance. Investigation of the compo- mortality rates, differential migration or spatial segre-
sition of bycatch of prawn trawlers on the east coast of gation by sex (Sadovy & Shapiro, 1987). There is
South Africa therefore provided an opportunity to evidence for disparity in growth rates for males and
describe aspects of the biology of four little-studied females of O. ruber and A. nibe, with females growing
sciaenids, Otolithes ruber, Atrobucca nibe, Johnius dus- faster in both species and also attaining larger sizes
sumieri and J. amblycephalus. Sciaenids are often the (Fennessy, unpubl. data). The observed skewed sex
dominant taxon in turbid coastal waters off river ratios in larger size classes of A. nibe (Figure 3) may
mouths (Lowe-McConnell, 1977; Trewavas, 1977; also have been caused by spatial segregation of the
Yanez-Arancibia et al., 1985; Longhurst & Pauly, sexes, as deep-water trawl catches consisted almost
1987), and have been reported as a bycatch of prawn entirely of females of this species (pers. obs.).
trawlers in several countries (Saila, 1983; Brinca et al., The extended breeding seasons observed in all four
1984; Sheridan et al., 1984; Blaber et al., 1990). species appear to be typical of sciaenids in subtropical
The skewness of the length frequency distributions waters (Druzhinin, 1974). Spawning in spring and
of O. ruber, A. nibe and J. amblycephalus caught by summer has been reported for other sciaenids in
shallow-water prawn trawlers (Figure 2) indicates that South Africa, specifically A. japonicus (Griffiths,
either larger fish are not as vulnerable to the gear, or 1996a) and A. aequidens (Griffiths & Hecht, 1995a),
they are not common on the Tugela Bank. However, although in the warmer east coast waters, spawning by
the same modus operandi is employed by the deep- these species commences in winter. Previous studies
water trawlers, which catch A. nibe that are consider- have reported spawning of O. ruber in January and
ably larger than those caught by Tugela Bank trawlers February on the KwaZulu-Natal coast, and from
(Figure 2). This implies that the gear is sufficiently September to February for J. dussumieri (Wallace &
effective to catch larger fish, and hence, the likely van der Elst, 1975).
explanation for the lower numbers of larger individ- Few previous estimates of size at maturity are
uals of O. ruber, A. nibe and J. amblycephalus on the available for the species investigated here. Wallace
Tugela Bank is that they are not numerous there. (1975) estimated a size at 50%-maturity (both sexes
The deviations from a 1:1 sex ratio in the larger size combined) of 10–12 cm (total length, TL) for J.
classes of all four species suggest sex change (Wenner, dussumieri, and 22–24 cm TL for O. ruber on the
264 S. T. Fennessy

(a) mated size at 50%-maturity for A. nibe to be 145 mm


2 17 107 208 198 150 117 136 146 143 144 80 33 18 4 3 TL. Apart from this last estimate, values from the
100
literature agree closely with estimates obtained in this
75
study.
%

50
25 The diets of the four species caught in trawls
0 consisted mostly of small epibenthic crustaceans, par-
105 125 150 175 200 225 250 275 300 325 350 375 400 425 450 475
ticularly penaeid prawns and brachyurans, and small
TL (mm)
(b) fish, all of which were commonly recorded in the
1 17 39 30 2 3 14 99 152 162 122 146 94 24 11 2 trawler bycatch (Fennessy et al., 1994; Fennessy,
100 1995). Several other studies have also demonstrated
75
the importance of crustaceans in the diets of these
%

50
25
and other sciaenids (Valdya, 1960; Savant, 1963;
0 Druzhinin, 1974; Sato, 1974; Brinca et al., 1983;
125 150 175 200 225 250 275 300 325 350 375 400 425 450 475 500
Pillai, 1983; Passoupathy & Natarajan, 1987).
TL (mm) The information on length frequencies, sizes at
(c)
1 23 59 100 165 279 286 224 133 70 20 2
maturity and feeding presented here points to the
100 functioning of the Tugela Bank as a nursery area for
75 O. ruber, A. nibe and J. amblycephalus, and, to a lesser
%

50
extent, J. dussumieri. Most trawled individuals of
25
the first three species are immature (Figure 2), and the
0
60 75 90 105 120 135 150 165 180 195 210 225 majority of adults appear to be located elsewhere. The
TL (mm) size and frequency of O. ruber caught by boat anglers
(d)
off Durban (Figure 2) indicates that there is dispersal
51 169 190 163 93 127 92 108 111 102 44 4
100 of the adults of this species to other areas of the shelf.
75 In the case of A. nibe, the adults occur on the
%

50 continental shelf break, where they are caught by


25 deep-water prawn trawlers (Figure 2). Very little
0
90 105 120 135 150 165 180 195 210 225 240 255 sampling of fishes has occurred on the shelf in inter-
TL (mm) mediate depths i.e. between 50 and 350 m, so it is
F 3. Sex ratios of four sciaenid species trawled on the possible that the adults also occur there. There is
KwaZulu-Natal coast. Solid bars=female, clear bars=male. some evidence for this, as there is an increase in the
Sample sizes for each size class indicated on top x-axis; size mean sizes of all four species with increasing depth on
classes which differed significantly from 1:1 have the sample the Tugela Bank (Table 2). The utilization by J.
size enclosed in a box (P=0·05). (a) O. ruber; (b) A. nibe;
(c) J. dussumieri; (d) J. amblycephalus. dussumieri of the Tugela Bank as a nursery is less
apparent, as most of the length classes are well repre-
sented (Figure 2). This euryhaline species is classified
T 4. Length-weight equations for four sciaenid species as partially estuarine-dependent (Whitfield, 1994),
from KwaZulu-Natal (May 1989 to February 1994) with some juveniles utilizing estuaries as nurseries
(Wallace & van der Elst, 1975). This probably
Species L/W equation R2 n
accounts for the reduced abundance of smaller,
O. ruber W=4·94E-06(L)3·13 0·996 2511
immature size classes in Tugela Bank catches relative
J. dussumieri W=3·08E-06(L)3·28 0·984 1782 to the other three species. Based on Figure 2, how-
J. amblycephalus W=3·43E-06(L)3·25 0·989 1544 ever, a large proportion of immature J. dussumieri also
A. nibe W=4·58E-06(L)3·15 0·995 1437 occur offshore. Further evidence for the nursery func-
tion of the Tugela Bank for sciaenids is revealed by the
results of the stomach content analysis. The occur-
KwaZulu-Natal coast (versus 125 mm and 237 mm rence of small-sized, abundant prey in the area
in this study, respectively). The former estimate was (Fennessy, 1995) undoubtedly contributes to the suit-
based on estuarine specimens. Vaidya (1960) esti- ability of the Tugela Bank as a nursery area. The
mated size at first maturity for female O. ruber to functioning of this area as a nursery has also been
be 220 mm TL, and Sato (1974) estimated a size suggested for another sciaenid Argyrosomus thorpei
at maturity (combined sexes) for Argyrosomus (Fennessy, 1994).
(Atrobucca) nibe from Japan of 24 cm TL, versus There are very few data available to indicate that
286 mm in this study. Murty (1980), however, esti- other areas of the KwaZulu-Natal coast or shelf serve
Biology of Sciaenidae from South Africa 265

(a) (b)
7 11
n = 277 n = 436
10
6
9

5 8
Gonadosomatic index

Gonadosomatic index
7
4
6

3 5

4
2
3

1 2

1
0
n
b
ar
pr
ay

n
l
ug

p
ct
ov
ec

n
b
ar
pr
ay

n
l
ug

p
ct
ov
ec
Ju

Ju
Fe

Fe
Se

Se
Ja

Ju

Ja

Ju
O

O
D

D
M

M
A

A
N

N
M

M
A

A
(c) (d)
8 9
n = 166 n = 188
7 8

7
6
Gonadosomatic index

Gonadosomatic index

6
5
5
4
4
3
3
2
2

1 1

0 0
n
b
ar
pr
ay

n
l
ug

p
ct
ov
ec

n
b
ar
pr
ay

n
l
ug

p
ct
ov
ec
Ju

Ju
Fe

Fe
Se

Se
Ja

Ju

Ja

Ju
O

O
D

D
M

M
A

A
N

N
M

M
A

F 4. Gonadosomatic indices of females of four sciaenid species from KwaZulu-Natal. Dashed lines means missing data.
Bars indicate one standard deviation. (a) O. ruber; (b) A nibe; (c) J. amblycephalus; (d) J. dussumieri.

as nurseries for the four sciaenids investigated here. sciaenids are apparently low elsewhere along the
The dependence of juvenile J. dussumieri on estuaries coast. Finally, Beckley and Fennessy (1996) and
has already been mentioned, but it is noteworthy that, Fennessy (unpubl. data) found few juvenile individ-
of all the estuaries on the KwaZulu-Natal coast, it is uals of O. ruber, J. dussunieri and J. amblycephalus from
only in the St Lucia estuarine system that this species sandy substrates in the Durban area.
appears to occur in significant numbers (Begg, 1984). Several authors have identified shallow marine
Further, Wallace (1975) notes that, in the St Lucia (non-estuarine) areas as nurseries for a variety of
system, adults of this species far outnumber juveniles. fish species (e.g. Beckley, 1984; Buxton et al., 1984;
Despite high angling effort levels in KwaZulu-Natal, Smale, 1984; Bennett, 1989). Although some
catches from the shore and from offshore boats sciaenids do utilize estuaries for this, several species
show only very low numbers of O. ruber (Mann et al., make use of inshore waters (Lowe-McConnell,
1997), with the implication that abundances of small 1966; Lowe-McConnell, 1977; NOAA, 1985). Blaber
266 S. T. Fennessy

(a)
100
80

No trawl data
60

%
40
20
0
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

(b)
100
80
60
%

40
20
0
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

(c)
100
80

No trawl data
60
%

40
20
0
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

(d)
100
80
No trawl data

60
%

40
20
0
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec
Inactive Active Ripe Spent
F 5. Proportions of gonad maturity stages of females of four sciaenid species from KwaZulu-Natal. *Indicates the
occurrence of partly-spawned fishes in samples. (a) O. ruber; (b) A. nibe; (c) J. dussumieri; (d) J. amblycephalus. Note that there
are no trawl data for three species in December.

et al. (1995), specifically noted that O. ruber and suitable nursery areas for fishes i.e. food, turbidity and
J. amblycephalus utilized the latter as a nursery. Blaber shelter (Blaber & Blaber, 1980). The occurrence of a
and Blaber (1980); Lenanton (1982); Blaber et al. rich food supply on the Tugela Bank has already been
(1989) and Ansari et al. (1995), amongst others, referred to previously. The turbidity which character-
particularly specified the importance of turbid, coastal izes the sea in the area is a result of the numerous
marine areas as nurseries. Blaber (1981) maintains rivers which flow into the sea from the adjacent coast,
that, unlike south-east Asia, such areas are lacking in which in turn have contributed to the development of
south-east Africa, and suggests that estuaries fulfill the the only extensive depocentre of mud on the east coast
function of these habitats on the South African coast. of South Africa (Flemming & Hay, 1988). Finally, the
However, the Tugela Bank possesses some of the Tugela Bank is situated within the Natal Bight, which
qualities which characterize estuaries and make them forms an indentation in an otherwise linear coastline
Biology of Sciaenidae from South Africa 267

(a) (b)
100 100
n = 397 n = 709

80 80

Percent mature
Percent mature

60 60

40 40

20 20

0 100 200 300 400 500 0 100 200 300 400 500
Total length (mm) Total length (mm)

(c) (d)
100 100
n = 343 n = 310

80 80
Percent mature

Percent mature

60 60

40 40

20 20

0 50 100 150 200 250 0 50 100 150 200 250


Total length (mm) Total length (mm)
F 6. Plots of sizes of 50%-maturity of females of four sciaenid species from KwaZulu-Natal. (a) O. ruber; (b) A. nibe;
(c) J. amblycephalus; (d) J. dussumieri.

(Figure 1). The continental shelf is at its widest here of protection from the rough seas which otherwise
(Lutjeharms et al., 1989), so there is some degree characterise the east coast. These factors combine to
produce a suitable nursery area for the sciaenid
species investigated here.
T 5. Estimates of size-at-50-% maturity for females of
four sciaenid species from KwaZulu-Natal (May 1989 to
February 1994) derived from the fitting of logistic curves Acknowledgements
to maturity data. m=estimates of the mean
This study was funded by the South African
Size at 50% 95% confidence
Species maturity (mm) interval R 2 Association for Marine Biological Research and the
Sea Fisheries Fund. Thanks are due to the Director
and staff of Natal Ocean Trawling for facilitating the
O. ruber 237 236cmd239 0·998
collection of samples and to the Natal Sharks Board
A. nibe 286 270cmd302 0·895
J. dussumieri 125 120cmd131 0·940 for logistical support. Three anonymous referees and
J. amblycephalus 168 162cmd174 0·965 colleagues at the Oceanographic Research Institute
provided useful comments on the manuscript.
268 S. T. Fennessy

T 6. Diets of four Tugela Bank sciaenids (May 1989 to February 1994), represented as number
of stomachs

Class Taxon O. ruber J. dussumieri J. amblycephalus A. nibe

Polychaeta 1 1 21
Crustacea Copepoda 2
Ostracoda 13
Mysidacea 4 5 1
Stomatopoda 3 9 13
Natantia 356 139 135 18
Anomura 3 30 38 3
Brachyuran 88 24 39
Cephalopoda 9 1 1 2
Pelecypoda 2 1 3
Gastropoda 1
Asteroidea 1
Osteichthyes 117 55 14 55
Vegetation 1
Digested matter 37 106 96 2
Empty 1092 933 882 604
Total 1713 1319 1244 685

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