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Volume 51, 2000
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 Mar. Freshwater Res., 2000, 51, 777–82

Occurrence of tuna and mackerel larvae (Family: Scombridae)

off the east coast of South Africa

Lynnath E. BeckleyA and Jeff M. LeisB

A
Oceanographic Research Institute, PO Box 10712, Marine Parade, Durban 4056, South Africa
email: seaworld@dbn.lia.net
B
Australian Museum, 6 College Street, Sydney, NSW 2010, Australia

Abstract. Spatial and temporal distribution patterns of scombrid larvae along the east coast of South Africa were
investigated from ichthyoplankton collections made during May–June 1990 (winter), October 1990 (spring) and
February 1991 (summer). Results were analysed in relation to oceanographic conditions and known spawning local-
ities of tuna and mackerels in the western Indian Ocean. In total, eight species were represented in the samples, with
highest diversity in February and lowest numbers in May–June. Larvae of the temperate chub mackerel Scomber
japonicus were most abundant at shelf stations during October. Larvae of neritic tunas Auxis sp. and Euthynnus
affinis occurred in shelf stations off KwaZulu–Natal in February and extended southward in a plume along the shelf
edge. Larvae of skipjack tuna Katsuwonus pelamis were most abundant in the Agulhas Current during February.
Only a few larvae of oceanic tunas Thunnus spp., wahoo Acanthocybium solandri and king mackerel
Scomberomorus commerson were collected in the Agulhas Current in the north of the study area during February
when there was an intrusion of warm Tropical Surface Water. This indicates that spawning of these species probably
does not occur off the east coast of South Africa.

Extra keywords: Scomber japonicus, Auxis sp., Euthynnus affinis, Katsuwonus pelamis, Thunnus spp.

Introduction
The pelagic teleost family Scombridae is well represented in
South African waters, and several species of tunas and mack-
erels contribute to the country’s commercial and recreational
fisheries. For example, there is a long-established pelagic
seine fishery for chub mackerel Scomber japonicus in the
Western Cape (Crawford 1989) and, since the early 1980s,
albacore Thunnus alalunga, in particular, has been fished in
the south-east Atlantic Exclusive Economic Zone (EEZ) by
the poling method (Penney and Punt 1993). Along the east
coast there is an extensive boat-based recreational fishery,
and species such as king mackerel Scomberomorus commer-
son, skipjack Katsuwonus pelamis and yellowfin tuna
Thunnus albacares form a significant part of the catch, par-
ticularly off the KwaZulu–Natal coast (Penney et al. 1999).
Despite the existence of these fisheries, little attention has
been paid to the biology – in particular the reproductive
biology and the early life history stages – of scombrid fishes
in South African waters.
The east coast of South Africa is characterized by the
Agulhas Current, a powerful western boundary current that
flows south-westwards along the edge of the narrow conti-
nental shelf getting further from the coast as the shelf widens
over the Agulhas Bank until it retroflects and heads east-
wards. The upper layers of the Agulhas Current are a mixture
of both Tropical Surface Water and Subtropical Surface
Water (Shannon 1989). During the course of ichthyoplankton
surveys along the east coast in 1990/91 to investigate the pos-
tulated role of the Agulhas Current in the dispersal of the
early life-history stages of fishes caught in the inshore hook
and line fishery (Beckley 1993, 1998) numerous scombrid
larvae were collected. In this paper, information on their
abundance and distribution along the east coast is presented
and discussed relative to oceanographic conditions and
known spawning locations of scombrids in the south-western
Indian Ocean.
Materials and methods
Three cruises were conducted with the RS Sardinops along the east
coast of South Africa in May–June 1990, October 1990 and February
1991. The cruises covered a grid of stations extending from Algoa Bay
(34°S) to Tugela (29°S), a longshore distance of ~800 km (Fig. 1). Nine
station lines perpendicular to the coast were occupied. Each line had two
stations on the shelf in depths of 50 m and 100 m and two stations in the
Agulhas Current offshore of the shelf break in depths of 500 m (slope)
and 2000 m (oceanic). The offshore extent of the grid varied from 30 km
to 100 km depending on the width of the continental shelf and adjacent
bathymetry. The outer station on the Tugela line was restricted to 1000
m depth because Grundlingh (1983) had shown that the mean position
of the core of the Agulhas Current was inshore of the 1000 m isobath

© CSIRO 2000 10.1071/MF00044 1323-1650/00/080777

778
Fig. 1. Sampling stations and sea-surface temperature isotherms during three cruises to collect ichthy-
oplankton off the east coast of South Africa in 1990/91.
along this part of the coast. The 2000 m stations on the Durban and Port
Alfred lines could not be sampled during June 1990 and October 1990,
respectively, because of severe gales.
Oceanographic conditions were recorded during the three cruises
and the results were described in detail by Beckley and van Ballegooyen
(1992). Sea-surface isotherms are included in Fig. 1 to give some indi-
cation of prevailing conditions. Ichthyoplankton samples were col-
lected with standard bongo nets of 57 cm mouth diameter and 500 µm
mesh equipped with mechanical flow meters and a recording diving
depth gauge. Stepped oblique tows were conducted in the upper 80 m of
the water column at all stations except at the inshore 50 m depth stations
where only the upper 40 m was sampled. All tows were of 10 minutes dura-
tion, and mean volume of water filtered per net was 197 m3 (s.e. 2.5 m3).
This methodology, with a slower net retrieval rate for the inshore 50 m
depth stations, can result in some bias towards these stations if particular
larvae only occur in the upper part of the water column.
Samples were preserved in 5% v/v formaldehyde in sea-water. Fish
larvae were separated from the zooplankton under a dissecting micro-
scope. Scombrid larvae were identified by reference to published
descriptions (Collette et al. 1984; Jenkins et al. 1984; Nishikawa and
Rimmer 1987; Richards 1989; Leis and Trnski 1989; Collette and
Aadland 1996). For abundance and distribution information, data from
both sides of the bongo net were combined at each station.
Results
Of the larvae collected, 3992 (nearly 13%) were scombrids;
these represented eight species, and highest diversity
occurred during the February 1991 cruise (Table 1).
Lynnath E. Beckley and Jeff M. Leis
Larvae of the temperate chub mackerel Scomber japoni-
cus were extremely abundant in the spring of 1990, with 3425
sampled at stations on the continental shelf and shelf edge
throughout the study area during the October cruise.
Densities in excess of 50 S. japonicus larvae per 100 m3 were
recorded at eight stations, with a maximum density of 205
larvae per 100 m3 at the 50 m station off Durban (Fig. 2).
Many of these larvae were still in the yolk-sac stage, and
highest densities of these newly hatched larvae were found at
the 50 m stations off East London and Green Point, at the 50
m and 100 m stations off Durban, and at the 500 m station on
the shelf-edge of Algoa Bay.
Larvae of the other scombrid species were recorded
mainly during the summer cruise in February 1991. The
larvae of the neritic frigate tuna Auxis sp. were the most abun-
dant and occurred in a plume extending southwards from the
KwaZulu–Natal shelf along the edge of the Agulhas Current
(Fig. 2). Maximum densities were in the region of 15–20
larvae per 100 m3 and occurred at the shelf stations off
Durban and Green Point. Larvae of eastern little tuna
Euthynnus affinis were less abundant but at this time also
occurred in shelf waters along the KwaZulu–Natal coast and
in a plume extending southwards (Fig. 2).
Few larval specimens of the wide-ranging oceanic tunas
were recorded. Only two larvae of the skipjack tuna
Occurrence of tuna and mackerel larvae off South Africa 779

Table 1. Abundance of scombrid larvae during three cruises off the
east coast of South Africa in 1990/91
Species
Acanthocybium solandri
Auxis sp.
Euthynnus affinis
Katsuwonus pelamis
Scomber japonicus
Scomberomorus commerson
Thunnus alalunga
Thunnus albacares
Thunnus sp.
Unidentified scombrid
Total
Katsuwonus pelamis were recorded in each of the May/June
1990 and October 1990 cruises, but 40 specimens were
obtained from samples taken in the Agulhas Current during
May/June October February
the February 1991 cruise (Fig. 2). The three albacore
1990 1990 1991
Thunnus alalunga larvae recorded were collected in
4 February 1991 in the Agulhas Current at the northern extreme
2 466 of the study area at the outer station of the Tugela, Durban and
1 36 Green Point lines. A single yellowfin tuna Thunnus albacares
2 2 40
larva was recorded at the same time at the outer station on the
3 3425 1
1 Tugela line. Three other unidentified Thunnus sp. larvae were
3 also recorded at the Agulhas Current stations off Durban and
1 Green Point during the February 1991 cruise.
3 The February 1991 cruise also yielded four larvae of
2
wahoo Acanthocybium solandri, one at each of the outer sta-
5 3432 555 tions on the Tugela, Green Point and Port Edward lines as
well as one at the 100 m depth station on the Tugela line.

Fig. 2. Distribution and abundance of larvae of Scomber japonicus, Katsuwonus pelamis, Auxis sp. and Euthynnus affinis off the east coast
of South Africa in 1990/91. Dots indicate stations where larvae of that species were recorded.
780
A single larval specimen of the king mackerel Scomber-
omorus commerson was also collected on this cruise at the
100 m depth station off Port Edward.
Discussion
In general, the results of the ichthyoplankton survey have
indicated that distinct assemblages of larvae occur off the
east coast of South Africa at different times of the year and
these appear to be linked to seasonal oceanographic condi-
tions (Beckley 1998; Beckley and Hewitson 1994; Olivar and
Beckley 1994). The scombrid larvae showed clear spatial and
temporal variation in distribution and abundance.
The occurrence of larvae of the temperate chub mackerel
S. japonicus during the spring cruise was unexpected because
this species has previously only been recorded as spawning in
the upwelling areas off the south-western Cape with a peak
during June–September (Baird 1977). The presence of yolk-
sac-stage larvae strongly suggests that this species spawns
along the east coast. During the October 1990 cruise, temper-
ature of the shelf waters was coolest (Beckley and van
Ballegooyen 1992) and, coincidentally, at this time larvae of
the temperate sardine Sardinops sagax also reached
maximum abundance (Beckley and Hewitson 1994).
The summer cruise in February 1991 yielded the highest
diversity of scombrid larvae, which comprised both neritic
and oceanic species. Oceanographic conditions at this time
revealed a strong seasonal input of 28°C Tropical Surface
Water into the Agulhas Current in the north of the study area
and warm water of ~24°C on the shelf off KwaZulu–Natal.
Larvae of the two neritic species Auxis sp. and E. affinis were
most abundant on the shelf off KwaZulu–Natal with a plume
of larvae extending southwards. The larvae included many
preflexion specimens, indicating that spawning could have
occurred in the region. Conand and Richards (1982) recorded
Auxis sp. larvae in the Mozambique Channel and north of
Madagascar and E. affinis larvae off the west coast of
Madagascar and off the coast of Mozambique, and
Nishikawa et al. (1985) indicated that Auxis and Euthynnus
larvae extend south to 32°S in the western Indian Ocean.
K. pelamis larvae were also most abundant during the
summer cruise and this agrees with the observations of
Conand and Richards (1982). They recorded skipjack larvae
in the waters to the north and west of Madagascar extending
across to the East African coast and noted that abundance of
skipjack larvae was particularly high in the northern
Mozambique Channel during summer. They postulated that
there was summer spawning as far south as 30°S because of
the seasonal southward extension of the warm water (>27°C)
favoured by skipjack larvae. Nishikawa et al. (1985) recorded
scattered occurrences of K. pelamis larvae south to 28°S in the
western Indian Ocean. Stequert and Marsac (1989), in their
review of tuna fisheries in the Indian Ocean, mentioned that
skipjack larvae are found in most parts of the Indian Ocean,
from 36°S on the western side to 30°S on the eastern side, and
Lynnath E. Beckley and Jeff M. Leis
their accompanying map based on the early work of Ueyanagi
(1969) showed a clear plume of larvae extending south in the
region of the Agulhas Current. Thus, although the presence of
K. pelamis larvae in the study area might reflect spawning in
the extreme north-east of the South African EEZ, advection by
the powerful Agulhas Current of larvae spawned in the
Mozambique Channel is most probable.
The limited number of Thunnus larvae collected off the
east coast suggests that there is no major spawning area for
these oceanic tunas off the South African coast. Ueyanagi
(1969) indicated that spawning of these two species occurred
to the east of Madagascar during the summer months.
Nishikawa et al. (1985) noted the occurrence of T. albacares
larvae in the Mozambique Channel from 20°–25°S. Conand
and Richards (1982) frequently observed T. albacares larvae
to the north-east of Madagascar during the summer but found
T. alalunga larvae to be scarce, with the only specimens
caught being at 23–24°S east of Madagascar. Stequert and
Marsac (1989) concluded that albacore larvae are distributed
in two separate and distinct zones in the Indian Ocean, one
east of Madagascar and the other off Western Australia.
Yellowfin tuna larvae, however, occur more widely with
greatest larval concentrations south of Indonesia, east of
Madagascar and around the Maldives, Chagos and
Seychelles islands. Davis et al. (1990) investigated the verti-
cal distribution of T. albacares, T. maccoyii and K. pelamis in
the East Indian Ocean north-west of Australia and found
larvae of these oceanic tunas to occur above the pycnocline.
In the present study, the upper mixed layer in the Agulhas
Current rarely extended below 70 m depth (Beckley and van
Ballegooyen 1992), so by sampling to 80 m depth it is
unlikely that larvae of oceanic tunas were missed.
The presence of four A. solandri larvae during the summer
cruise suggests that, like the larvae of oceanic tunas, they
were advected into the region by the Agulhas Current.
Conand and Richards (1982) recorded a few A. solandri
larvae off the Comores Islands, and van der Elst (1981) stated
that spawning of this species has been recorded off the coast
of East Africa during most months of the year.
Only a single king mackerel (S. commerson) larva was col-
lected during the study and no larvae of the queen mackerel
S. plurilineatus were recorded. Harris et al. (1999), who
investigated the larval fish assemblage in the nearshore waters
~100 km north of Tugela, recorded 12 Scomberomorus sp.
larvae during monthly sampling. Adults of the two
Scomberomorus species are seasonally abundant off the
KwaZulu–Natal coast, and a detailed study of the biology of
S. plurilineatus by van der Elst and Collette (1984) revealed a
high incidence of fish in post-spawning condition leading
them to suggest that this species spawns during late winter and
spring off the coast of Mozambique. Ripe king mackerel have
been recorded in southern Mozambique during spring and
this, together with the low incidence of such specimens from
South Africa, (Oceanographic Research Institute, unpub-
Occurrence of tuna and mackerel larvae off South Africa
lished data) also suggests spawning to the north along the
Mozambique coast for this species. Jenkins et al. (1985) have
shown that the larvae of Scomberomorus spp. are coastal in
distribution and the possibility does exist that larvae in coastal
waters could have been missed during the present study
because all samples were taken in water >50m depth.
Several ichthyoplankton studies off the east coast of
Australia have referred to scombrids, but most have dealt
with tropical areas such as the Great Barrier Reef (e.g. Leis
and Goldman 1984; Jenkins et al. 1985; Thorrold 1993). Off
the more temperate New South Wales coast, Gray (1993) has
recorded an increase in abundance of larvae of unspecified
Scombridae in spring.
There are several similarities between the findings of the
present study and those of Matsuura and Sato (1981) who
examined the distribution and abundance of scombrid larvae
in southern Brazilian waters. In their study, scombrid larvae
were also least abundant in the autumn/winter and they also
found high concentrations of S. japonicus during spring. Of
the tunas, Auxis sp. and Euthynnus alletteratus larvae were
the most abundant, occurring in late spring and summer. Very
few larvae of K. pelamis and Thunnus sp. were collected and
those were from offshore stations in the Brazil Current. In
contrast, studies around tropical oceanic islands have
revealed high concentrations of tuna larvae in near-reef
waters (Leis et al. 1991; Boehlert and Mundy 1994).
In general, the findings of the surveys off the east coast of
South Africa concur with results from other Southern
Hemisphere studies and highlight the low abundance of
scombrid larvae in winter, the spring abundance of larvae of
the more temperate species S. japonicus, the summer occur-
rence of larvae of various tuna species and the role of western
boundary currents in distributing oceanic tuna larvae south-
wards from more tropical areas. The implication of this infor-
mation is that South Africa has to ensure the sustainable use
of S. japonicus because the life cycle is completed in the
South African EEZ but, for other scombrids, the fate of the
stocks is in the hands of those countries with which the
resource is shared, as well as requiring sound management by
the Indian Ocean Tuna Commission.
Acknowledgements
Thanks are extended to the master, officers and crew of the RS
Sardinops for their assistance, Cameron Barnard, Lynne
Shannon and Audrey Hutchings who sorted the fish larvae
from the plankton samples and Bernadine Everett who assisted
in preparation of the figures. The project was funded by the Sea
Fisheries Research Fund, South African Association for
Marine Biological Research, Foundation for Research
Development and University of Natal Research Fund.
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Manuscript received 24 March 2000; revised and accepted 24 May 2000