Marine fish eggs and larvae from the east coast of South Africa

Allan Connell

Durban

South Africa

Originally posted May 2007; updated October 2012.

Abstract. Twentyfive years of records are summarised, in 226 separate species data sheets, of eggs and early larvae of fishes spawning pelagic eggs on the inshore shelf, within 5km of the coast, along a short section of the KwaZulu-Natal coastline, about 50km south of Durban. Annual spawning period, and egg abundance trends over the 25 years, are provided, as well as egg and larval descriptions, showing the ephemeral pigment patterns of many species' early larvae, in digital image colour. By collecting both offshore (5km) and inshore (0.5km) samples, comparing percentage representation of each species within these two sample sets, and using two indicator species, the kob Argyrosomus japonicus and the geelbek Atractoscion aequidens, both with well-defined spawning grounds, a reasonable assessment of location of spawning was obtained for all the common eggs in the study area. Roughly 65 species were reared on a simple food-chain for identification purposes, but in the latter stages of the study, larval identification was by DNA barcoding using the cytochrome C1 gene. The study is ongoing, both to increase the annual trend graphics for each species, and to gather barcodes of currently unidentified eggs and larvae, so that they will be identified when adult material has been sequenced, in support of which, over 900 species of local marine fishes have been barcoded. Introductory Notes describe the geography and oceanography of the study area, with particular emphasis on how they affect spawning migrations of species into the study area, and movement of early juveniles to their preferred nursery areas.

1.                 Introduction

This study resulted from concern regarding the effects of effluent discharges into the marine environment, on the early life stages of marine fishes with pelagic eggs. In 1985, a major effluent pipeline was about to begin discharging industrial effluent over a shallow continental shelf area, offshore from the newly developed Richards Bay harbour, on the east coast of South Africa, in an area of widened shelf waters which was regarded as biologically productive in the local context (Figures 1 & 2). In an effort to assess whether the area was an important spawning ground for coastal marine fishes, a series of surface plankton samples was collected, over several years, to assess the diversity of fish species spawning in the area, and the intensity and seasonality of spawning. These samples were preserved with formalin, aboard ship, in order that they could be returned to the laboratory for further study.

Figure 1
Figure 1: Overview of the study area, on the east coast of South Africa

The samples soon highlighted the difficulty of trying to identify fish eggs fixed in formalin, without the benefit of being able to hatch the eggs, and examine the pigment patterns and other diagnostic features of the developing larva. Thus, while the Richards Bay study continued to examine formalin-fixed samples, a second study was started, closer to home. This entailed collecting live plankton samples, using a surface-towed plankton net, which enabled me to return the samples, alive, in 25 litre buckets of seawater, to the laboratory for separation, cataloguing, and hatching of the eggs. This latter study began in 1986, and careful cataloguing of eggs began in January 1987.

During the course of the study, it became apparent that certain important species were excluded, as they were attracted to estuarine systems. Thus, over the period May 1990 to December 1994, a series of samples was collected from the entrance to Durban harbour. Some of these eggs were rarely or never encountered in the coastal samples, adding insight into the importance of Durban Harbour to estuarine fishes in the region, and yielding data on species that otherwise would not have been documented.

Many of the eggs encountered, could be identified from the literature, particularly when coupled with the clues contributed by the newly hatched larvae. A small, scattered, local literature, dating back to the “father” of marine biology in South Africa, JDF Gilchrist, was conveniently summarised in the pioneering research, in the regional context, of Brownell (1979). Other useful works included Delsman (1921-1938), Uchida et al (1958), a series of papers by Mito (1961-1963), Fahay (1983), Leis & Rennis (1983), Moser et al (1984), Okiyama (1988), Matarese et al (1989), Leis & Trnski (1989), Shao et al (2001), and most recently the huge and impressive work on Atlantic fishes, edited by Richards (2006).  But there were many eggs which could not be identified with any certainty. In the early stages of the study, hardy Pleuronectiformes, which could, literally, be reared in a glass bowl, gave incentive to attempt rearing of the unknown eggs, in the hopes of rearing at least one specimen to the point where fin counts and juvenile features aided in identification. Towards this end, a simple foodchain was set up, involving the green alga Chlorella, the rotifer Brachionus plicatilis and the brine shrimp Artemia.

In the latter stages of the project, it was becoming increasingly apparent that many important species were not amenable to being reared in the available facilities, with the available foodchain. While recognising that the answer probably involved DNA, most of the methods being used were not sufficiently rapid and economical for the purposes of this project.  In April 2003, an article appeared in New Scientist, (Hecht, 2003) outlining a technique, using a 650 base pair section of the Cytochrome C oxidase subunit 1, of mitochondrial DNA, to differentiate between cryptic species of butterflies and a variety of other insect species. An approach to this team of researchers, led by Prof. Paul Hebert at the University of Guelph in Canada, revealed advanced plans to launch an ambitious project to “barcode” the fishes of the world. Linking with the Barcode project served two important goals. Firstly, the fishes of South Africa would be barcoded, contributing to the world-wide effort, and secondly, this would provide the adult sequences that were essential to identify my larvae from their barcode sequences. In addition, the Barcode of Life Databank (BOLD) contains a much wider spread of adult sequences from around the world, and the collection is expanding rapidly. Further details can be accessed at http://www.barcodinglife.org.

2.       Methods

Except for occasional samples taken in the Durban area, and off Tongaat, the study has been conducted almost exclusively offshore of Park Rynie, some 60km south of Durban (Figure 2). This area was chosen because it has the most extensive reef in the Durban area, is only about 5km south of the Aliwal Shoal (an area which was being earmarked for special conservation), and has extensive sandy areas between the reefs that ensure a rich fish diversity. It also has a relatively safe and convenient ski-boat launch-site, although, being a surf launch out of a protected bay, remains strongly dependent on weather and sea conditions for safe launching (Plate 1).

Plate 1: The launch site at Park Rynie, on a good day.

2.1     Sampling Offshore

The net was an elongate cone, about 2 metres long, constructed of 300 micron aperture, monofilament netting, tapering to a detachable codend. The mouth was semi-circular in shape, with a diameter of 50 centimetres. Due to an 8.5cm elongation on each side of the arch, the mouth area was calculated as 1360cm2.  A small float was attached at each corner, and a lead weight at the middle of the arch, to ensure that the net settled in the water with the bar horizontal and at the surface. On each trip to sea the net was towed behind the boat at about 1-1.5knot for 10 minutes, at a site 4-5km offshore, where the water depth was 40-50metres. From early 1994, a second sample was collected on each outing, about 500m-800m offshore. During each trip, wind and current speed and direction were recorded. Notes were kept on conditions, if unusual, such as unseasonally cold water.

From time to time a calibrated, propeller-driven flowmeter (General Oceanic Model 2030R) was used to calculate the volume of water passing through the net.

2.2     Sampling in Durban Harbour Mouth

The net was pulled along a disused quay roughly 150 metres in length, on the north bank immediately east of the small craft harbour (Plate 2, white arrows). Due to the 3m high quay-wall, a 50m length of rope was used, and the net was hauled both ways along

      

Plate 2: Two white arrows show the quay where Durban Harbour mouth (DHM) samples were collected.

the quay, a distance of about 300m. On the KZN coast, the daytime spring high tide occurs at about 4pm local time, every two weeks. Studies have shown that these spring tide periods are peak spawning times for estuarine fishes (eg Garratt, 1998). But in order to combine sampling with dusk, which is around 17h30 in June (winter) and 19h00 in midsummer (late December), the guide used was to sample at dusk on the day on which the afternoon high tide coincided with sunset, using South African Navy tide- and sunset-tables. A single sample was collected every 2 weeks.

2.3     Recruitment sampling at the Lovu and Mkomazi estuaries

During 2005 the opportunity arose to sample larval fish recruitment to these two key estuaries in the area (Figure 2). This allowed comparison of  recruitment patterns with spawning patterns of such species as Monodactylus falciformis, Rhabdosargus holubi and mullet(LIIB9 & LIIIB7). The estuaries were sampled on afternoon incoming spring tides, targeting the inflow period as the flow became sufficient to support a net anchored in the shallow mouth. The net had a rectangular mouth of 0.8 x 1.2 metres, length of about 2 metres and mesh aperture of 500 microns. Samples were removed from the cod-end every 10 minutes. Three 10 minute samples were collected on each visit to each estuary. Due to the constraints of needing to sample the early tidal flow, the two estuaries were sampled on consecutive days. When the mouth of an estuary closed, the net was manually manoeuvred in the surf for 10 minute intervals (sea conditions permitting). Though not directly comparable, these surf samples served to show which species were present, and their relative numbers. Samples were preserved in formalin, for later counting and measurement.

2.4     Live egg sample handling in the laboratory

At sea, each sample was washed from the codend of the net into a 25 litre bucket, containing about 15 litres of seawater, and then sealed with a tight-fitting lid, for transport to the laboratory. In summer, buckets were kept cool with wet towels, to avoid accelerated egg development during transport. In the laboratory, the inshore sample was aerated, while the offshore sample was concentrated into a 500ml Pyrex bowl (12cm diameter and 6cm wall), and examined for eggs, with a dissecting microscope set on 6x magnification, and using reflected light. All eggs were removed to small bowls of clean seawater, using a range of small-bore pipettes. After searching the surface of the bowl, and especially its meniscus, the bottom of the bowl was also searched, as some eggs become less buoyant, and even sink, as they develop. All the eggs were then separated into “species” by external characteristics, and where uncertainty existed, or to check for correctness, specimens were placed in Pyrex bowls of clean seawater for hatching. It was usually possible to place at least three “species” in one bowl, by selecting obvious size or feature differences that ensured they could be recognised after hatching. The process was then repeated with the inshore sample. All this had to be completed on the evening of the day the eggs were collected, as most eggs hatched overnight. In cases where unusually high numbers of eggs were collected, the dominant species were ignored during separation. After separation of the less common eggs, the sample was fixed in formalin and the more common species counted by sub-sampling, at leisure, the following day.

A simple “key” based on the physical features of pelagic fish eggs, was used to separate eggs into basic groups. Unique alpha-numeric numbers were then assigned to each “species”, for cataloguing purposes. An extensive photo-album of eggs and early larvae was built up for each “species”, as larvae developed to the full extent allowed by the yolk and oil globule in the egg.

2.5     Larval rearing

Due to the time constraints of the initial egg sorting process, and since most sampling was done on weekends, a small laboratory was arranged at home, and the garden shed became a simple rearing facility. Thus the project became manageable, combining egg sampling with a personal passion for fishing and SCUBA diving.

A basic foodchain was developed, comprising a single celled green alga, Chlorella, cultured by the simple expedience of keeping a few tilapia, Oreochromis mossambicus, in 5000 litre plastic ponds of seawater. These were fed chicken pellets, and the water rapidly turned green. In separate smaller tanks of about 500 litres, the rotifer Brachionus plicatilis was maintained on a mixture of yeast cells, codliver oil, highly unsaturated fatty acids and Chlorella. When a suitable batch of eggs of an unknown species was encountered in a sample, a 50 litre rectangular glass tank was prepared by being half-filled with seawater, made green by the addition of Chlorella, and seeded with a screened and washed concentrate of Brachionus. The eggs or hatched larvae, were then added. Gentle aeration was supplied from a single stone at one end of the tank. A single fluorescent light, the same length as the tank, was housed in the lid, and provided 24-hour lighting. In summer these lids were left down as a temperature of about 25°C was needed, while in winter they were raised to allow cooling to 21°C. Using this method, by the time the larvae were ready to feed, usually about 5-7 days post-hatch, the rotifer population had enjoyed rapid growth and contained a good proportion of juveniles for the fish larvae to feed on. As the rotifers multiplied in the tank, they were thinned by in situ pouring of tank-water, dipped from within a 300µm mesh net, into a 50µm mesh net. Partial water replacement, and addition of Chlorella, both as food for the rotifer and for its antibiotic properties, was also done as necessary. Tanks were skimmed, at least daily, with a sheet of paper-towel, half laid on the water and drawn across the length of the tank, to remove oils and surface scum from the tank. Every attempt was made to collect growth series from successful rearings, as voucher specimens, for research purposes, to be accessioned into the SAIAB fishes collection at Rhodes University in Grahamstown.

2.6     Photography of eggs and larvae

All photography was done through a Zeiss binocular dissecting microscope, mostly at the maximum magnification of 40x, using 10x eyepieces. A single lens reflex camera, with the lens removed, and attached close up to the microscope eyepiece with a piece of plastic tubing and lens adaptor bayonet fitting, was light, quick and simple to use. Focussing was through the camera eyepiece and mirror. Three flashes were used, to avoid shake, one set off by the  button on the housing, the other two by slave, while the camera was briefly opened on B setting. All pigment patterns were found to be best observed using reflected light, with the eggs in small watch glasses. Both black and white backgrounds were useful, though black was preferred unless black pigment was being highlighted. Larvae were transferred to eyeglasses, using a wide-bore pipette, and then anaesthetised with a tiny crystal of MS222, before drawing the water level down, with a fine pipette, to induce the larva to lie on its side. For myomere highlighting in larvae, aligning the larva along a white and black edge (white sheet of paper under the bowl, on a black background) was helpful. This also worked well when looking for segmentation in the yolk, tinting in the oil globule and black pigments in the egg. Recently the system was converted to taking direct digital pictures, with the camera on Macro setting, and using a ring-light for all-round illumination. Simple plastic tubing attachments allowed photos to be taken as quickly as with the previous setup, an essential feature when working with anaesthetised specimens. Rapid irrigation with clean seawater revived all but the most sensitive specimens if the anaesthetic dose was not too strong.

2.7     DNA collection and analysis

For DNA sequencing, eggs were hatched as usual, and once larvae had fully pigmented eyes, they were anaesthetised with MS222, then photographed, prior to fixing in 98% alcohol. Individual larvae were placed in Matrix box vials, in 98% alcohol, in preparation for barcoding. Tissue samples from locally collected adult fish, were also sent for barcoding, to provide sequences that could be compared with hatched larvae sequences, thereby identifying eggs. Within BOLD, sequences of marine fishes from around the world provide further opportunities to find a sequence match, for species that have not yet been sequenced from local waters. The CO1 process is described in Hebert et. al. (2003). For more information on the barcoding initiative, visit http://www.barcodinglife.org.

3.       The Physical and Oceanographic Conditions in the study area

3.1     Climate and weather

An excellent review of the climate and weather patterns of the area can be found in Hunter (1988). What follows is a brief summary of those aspects considered relevant to the spawning study.

The climate of the KwaZulu-Natal coastal belt is generally described as humid subtropical with a warm summer. The coastline is fairly straight (Figures 1 and 2), with a slight slope, when viewed from north to south, of about 30° to the west.

3.2     Winds

The two dominant winds are from the northeast and southwest, and they are both essentially parallel to the coast. In the Durban to Park Rynie area, they occur about equally often in the summer (Figure 3). During winter, the south and southwest winds tend to be a little more frequent, and stronger (Figure 4). These southwest winds are associated with coastal lows (known locally as westerly busters), usually preceding a cold front, and the cold front itself. The generation of these coastal lows is well described by Hunter (1988).

The presence of the warm Agulhas Current offshore, and the rapid cooling of the hinterland at sunset, results in the land breeze becoming a nightly affair on quiet nights, especially during the winter months (Figure 4), when it is even strong enough to occlude the northeasterly during the night. Offshore this is also manifested as a dense bank of cloud over the Agulhas Current in winter. Hunter (1988) reported that the land breeze usually dissipates by 08h00, but in

Figure 3: Average wind conditions in December (from Hunter, 1988).

mid-winter, it often blows until 11h00 (A Connell, personal observation). The incidence of summer sea breezes, on the other hand, is reduced by the warm offshore current on the KZN coast (Preston-White 1969).

3.3     Rainfall

The annual rainfall along the coastal belt is between 1000 and 1100mm. It is a summer rainfall area, with above 70% falling in the summer months of November to March. Due to the high escarpment adjacent to the coast, most rivers have deeply cut courses, and the Mkomazi river, just north of the

Figure 4: Average wind conditions in June (from Hunter, 1988). But note that the wind rose for Durban has been replaced by one for Cape St Lucia (see Figure 2) because Durban Airport is acknowledged as poorly placed to adequately measure the incidence of offshore winds (Lundie 1979).

study area, extends to the mountainous escarpment some 150km from the coast. Although not a massive river, it has a MAR (mean annual runoff) of 1036x106 cu.m. High erosion rates in the hinterland cause frequent discolouration of coastal waters in the summer. Most of the shorter rivers have small coastal lagoons, confined between short, sand filled coastal floodplains and a sandbar across the mouth. They often only breach once or twice in the summer, or not at all during years of poor rains (Begg, 1978). During wet summers, tongues of silt-laden river water are a common feature, penetrating a kilometre or more into the sea adjacent to the mouth of the bigger rivers such as the Lovu (MAR 112x106cu.m.) and Mkomazi.

3.4            Coastal Oceanography

The KZN coast has a relatively narrow shelf (Figure 2), the 200m depth contour being within 20km of the coast for most of its length, and this is true of the Park Rynie area. Tide range is at most about 2m at peak spring tide, down to about 0.5m at neap tides.

3.4.1    The Agulhas current

The dominant oceanographic feature along the entire KZN coastline, is the Agulhas current, one of the major western boundary currents of the world. A review of the coastal oceanography of the area can be found in Schumann (1988). Water temperature in the current core, rises to about 28°C in summer, and drops to about 23°C in winter.

On average the Agulhas current is 40-60km offshore in the vicinity of Durban (Schumann 1988), but the inner edge can change its position by 30km and more in a day. Such meanders can be seen in Plates 4-8. Off Port Edward, Schumann (1988) reported that the Agulhas current was usually so close inshore that the inside edge, indicated by a sudden temperature rise of about 2°C, was seldom discernable,

3.4.2    Inshore waters and currents

 

Being a western boundary current, travelling south in the southern hemisphere, the Coreolis effect, also termed Ekman sheer, causes Agulhas current midwater to slide onto the shelf all year round (Pearce, 1978; Schumann 1988). This water is reported by Pearce (1978), to be from 40-60m depth in the Agulhas current, and is on average about 1.4°C cooler than Agulhas current surface water. In summer the surface of this cooler inshore water is rapidly warmed by the sun, resulting in a marked difference (about 4°C, according to Pearce, 1977), between surface and bottom temperature in shelf water 30-50m deep. Recent data from a thermister string deployed in 31m off Amanzimtoti demonstrates this (Figure 5). In winter, the weaker sun, along with the nightly cooling effect of the winter landbreeze, prevents this temperature difference from developing. Thus, in winter, inshore shelf water temperatures are generally within about 1°C from surface to bottom, at around 21°C (Figure 5). Countless satellite images

Figure 5

Figure 5.(Data courtesy J. Howlett, Huntsman Tioxide)

confirm the presence of cooler water inshore of the Agulhas current, as shown in the two examples in Plate 3 below.

Plate 3. Sea surface temperature images, summer (left), and winter (right), illustrating the presence of a band of cooler inshore water, relative to the Agulhas current, throughout the year along the KZN coast.

Plate4

Inshore currents are predominantly wind-driven, outside of the influence of topographically driven features such as the Durban-Richards Bay (A in Plate 4) and Port St Johns Bights (B in Plate 4). Although caused by relatively small kinks in the coastline, these are the sites of semi-permanent eddies of expanded cooler inshore water, with predominantly north-going currents on their inshore edges. Inshore currents in the Park Rynie area are largely wind driven, but complicated by southern extensions of the Durban Bight eddy, as is clearly demonstrated in Plate 4. This is confirmed by the unusual incidence of inshore currents off Park Rynie,  reported by Schumann (1988). The Durban Shelf area is described by Schumann, (1988) as “a transition region between the wind-dominated shelf to the north, and the Agulhas Current dominated shelf to the south”, with evidence

that the area between Durban and Park Rynie (his Mzinto) often shares the same water in a recirculating eddy.

A further disrupting feature to inshore currents, is the passage of a “Natal pulse”, essentially a moving eddy of water trapped inside the Agulhas Current. A “pulse” appears to be generated when a topographically generated eddy becomes unstable and breaks free to move down the coast (or a large inshore deflection of the Agulhas current becoming isolated), with a rapid clockwise rotation (when viewed from above), generating unusually powerful north-going currents on its inshore edge (Lutjeharms and Connell 1989). The big eddy off Hibberdene in Plate 7 is probably an example of this feature. A “pulse” moves down the coast at a speed of about 22km per day (Lutjeharms and Roberts 1988), and, due to their size

and depth, can be associated with cold water, and unusual, deepwater fauna being pulled up onto the shelf (eg squid egg balloons, and deepwater copepods). Most studies seem to suggest that these “pulses” are relatively infrequent, with only 3-4 passing through each year. The coldest water we ever encountered during the 25 years of diving related to this research, was 14.5°C, on 5 February 2006. That this was deep offshore water being pulled onto the shelf by a "Natal pulse", is supported by our collecting the first record of squid egg balloons from the east coast of South Africa (Roberts, Zemlak and Connell 2011). Such water would come from >150m over the shelf edge (Schumann 1988).

3.4.3  The Central Shelf

Plate 4 shows a band of cooler inshore water occupying the entire bight from Durban to just south of Richards Bay, caused by the Agulhas current being confined further offshore by the straight shelf break (Figure 2), with the coastline cutting away to the west, immediately south of Richards Bay. Flemming and Hay (1988) showed how bottom currents inferred from sediment dispersal and bedform patterns revealed a complex flow (Figure 6). These confirmed a closed eddy system off the Tugela mouth, and an accompanying depocentre for fine silts, as well as a semi-permanent (also cyclonic) eddy just north of Durban. In Plate 4 a similar eddy can be seen, centred just south of Port St Johns, extending north to about where 30°E longitude cuts the coast. For fishes moving up the coast on annual migrations, these two semi-permanent eddys form important “stepping stones”, since their inshore edges have predominantly north-going currents.

3.4.4  Coastal productivity, nutrients, zooplankton and fish

Text Box: Figure 6: Currents inferred from bedform patterns (from Flemming & Hay 1988)The Agulhas Current, being the dominant feature of the coastal oceanography of this region, also plays a major role in the year-long nutrient loading and productivity of the region. Studies have shown that surface water has a negative gradient in nutrient concentration with distance offshore as far as the Agulhas Current core [Oliff (1973), and Pearce (1977a) in Carter and D’Aubrey (1988)], as well as a positive gradient with depth in the Agulhas Current itself. The continuous Ekman veering of Agulhas Current midwater up onto the shelf, results in the inshore waters being cooler, and slightly elevated in inorganic nutrients, (N, P and Si), compared to the Agulhas Current core surface water. In KZN inshore waters, there is thus a strong correlation between the cooler inshore water, and chlorophyll-a production (Plate 9), and the chlorophyll-a levels are similar in summer and winter (Plate 10).

9A 9B

Plate 9. Demonstrating the strong relationship between the cool inshore water, and chlorophyll-a concentration, along the KwaZulu-Natal coast.

 

10A 10B

Plate 10. Comparison of chlorophyll-a concentration, in summer (left) and winter (right), along the KwaZulu-Natal coast.

 

Zooplankton has not been extensively studied on the KZN coast, and nothing has been added since the review of Carter and Schleyer (1988), apart from papers on fish larvae, which are not relevant to the present discussion. By far the dominant animals in most sea surface zooplankton samples are the copepods, and they are also the key component in the food web from phytoplankton to fishes. Transects across the Agulhas current have shown a higher biomass inshore of the current (Carter 1977), and the bulk of the biomass (70%) in the upper 100m (Carter and Schleyer 1988).   The dominant species in the core of the current was Paracalanus parvus, but on the shelf, Centropages chierchiae and Calanoides carinatus were dominant (Carter 1977). Carter (1977) recorded a shelf biomass minimum in the Durban area, in the late summer (March), and a winter biomass bloom dominated by the large calanoid, Calanoides carinatus. This is a key species in the diet of sardine; thus a winter bloom would be an important food source for this species during its annual winter migration to KZN waters. During the present study, notes were kept when big samples of large copepods were encountered in the egg samples, and by far the most common large copepod seen in winter, was Calanus agulhensis, also important in the diet of sardine.

Productive food chains, which in coastal seas lead to productive fisheries, are obviously dependent on sustained high primary production (phytoplankton), driven by nutrients, which are usually brought to the surface in cold upwelled water. This in turn supports a rich zooplankton, feeding on the phytoplankton bloom, and supporting fast-growing, pelagic, shoaling fishes such as sardines and anchovies. This is clearly not the case in KZN, due to the oligotrophic nature of the shelf waters, despite the Eckman-veer driven upwelling, and small areas of increased local upwelling such as the semi-permanent eddy located east of Durban, which can cause significant upwelling of nutrient richer, cold water at its centre (Pearce 1979), and the inshore waters south-west of Durnford Point (Lutjeharms et al 2000). The point is best illustrated by comparing the coastal fisheries of the South African coast. The KZN shelf does not support any year-round pelagic fishery, and only yields between 1500 and 2000 tonnes of linefish and about 500 tonnes of shoaling sardine-like pelagics, these latter associated with the annual winter “sardine-run” up the east coast from Cape waters (van der Elst 1988). By contrast, the southern Cape coast fishery yields in the region of 100 000 tonnes annually, and the western Cape fishery in the region of 600 000 tonnes annually. Without the zero’s the ratio is 2:100:600. Clearly the east coast is not a significant fishery. The real value of the KZN fishery, in the local context, is the quality of the fish harvested (van der Elst, 1988), which finds its way into local markets and onto local tables, via a small but active commercial fishery and an ever growing recreational fishery. Many of the fish species are of tropical origin; the warm, powerful Agulhas Current allows tropical species to extend their range further south than would otherwise be the case. But the area also exhibits a high degree of endemism, due again in no small measure to the powerful Agulhas Current which, along with the vast and open southern Indian Ocean, forms an effective barrier to coastal species migration, away from the southern tip of Africa.

4.       Coastal oceanography in relation to sardine movement into KZN waters in winter

The Agulhas Current plays a major role in shaping the so-called Natal sardine run each year. There is no evidence of a winter counter-current up the east coast, as has often been suggested in popular articles about the “Natal sardine run”. At all times of the year, shelf waters are a couple of degrees cooler than Agulhas current water. As winter approaches, inshore waters, continuously fed by Ekman veering of Agulhas current midwater (Section 3.4.2), gradually cool. As the sardine shoals move up the coast, in this cooler inshore water, they encounter, in the vicinity of Port St Johns, a narrowing of the shelf towards Port Edward (Plate 4), and consequently a narrowing of the band of cooler water between the Agulhas current and the coast. In this area there is evidence of an eddy formed in a similar way to the cooler water between Richards Bay and Durban (see Section 3.4.3), which ends in a bottleneck off Port Edward (Plates 4 - 8). While not consistently as wide as the Durban to Richards Bay complex of eddies, the slight westward slope of the coast south of Port Edward appears to contribute to this feature (Plates 3-10). Within this zone, the clockwise rotation of an eddy creates north-going inshore currents, which would assist the northward movement of the shoals. The Port Edward bottleneck, created by the very narrow shelf in that area (Schumann 1988) ensures that the shoals gather in the northern extremity of the eddy, ever closer to the coast. In some years the shoals get pushed through the bottleneck, by a severe cold front from the south-west, with strong southwesterly winds, spoiling the spectacle of the sardine shoals on the lower south coast of KZN. But in calmer years, the shoals will amass until the migratory imperative forces them to gather into a larger shoal which proceeds up the coast, hugging the coast to avoid the Agulhas Current offshore, in a long thin shoal that is often only about 50m wide, but can stretch 15-20km or more along the coast. If strong southerly currents are encountered just offshore, the “main” shoal can move in this fashion as far as Hibberdene or even Park Rynie, and has, occasionally, as on 30 June 2005 (Plate 11), even reached Durban in this mode.

plate11

Plate 11. Taken at Brighton Beach, Durban, on 30 June 2005, this image shows a thin continuous band of sardines (light purple in a blue sea), just behind the breaking wave. The shoal stretched for about 15km along the coast, at the time this picture was taken. The white dots and rafts in the background are seabirds (Cape Gannet), resting on the water after feeding on the sardines earlier in the morning.

Generally, however, they will break up into smaller shoals further down the coast, and continue moving north. Once under the influence of the predominantly north-going currents north of Durban, the sardines tend to disperse further offshore, where they feed and spawn in the cooler coastal waters inside the Agulhas Current, over the next few months (Connell 2001). As can be seen in Plate 4, another significant “bottleneck” is found north of Richards Bay, and although sardines have been found spawning off Richards Bay in October (Connell 1997a, 1998, 2003), there are no confirmed reports of shoals moving through the Cape St Lucia area. Connell (1997) showed, from evidence of egg collections off Park Rynie, that sardines move back south, through the Park Rynie area, from October to December each year, as KZN coastal waters become too warm. But due to the warmer surface waters inshore, they are not seen on the surface, and only the eggs indicate their presence as they move through the area (see notes, Sardinops sagax).

5.                 Larval fish movement in relation to the Agulhas Current.

A major feature of spawning patterns of fishes on the shelf of KZN, is that some of the key spawning species move into these waters, from the south, in winter, on spawning migrations (van der Elst 1988, Hutchings et al 2003). The most spectacular of these are the sardines, Sardinops sagax, the sciaenids Atractoscion aequidens (geelbek) and Argyrosomus japonicus (kob); the sparids Polysteganus undulosus (seventyfour), Petrus rupestrus (steenbras), Cymatoceps nasutus (poenskop), Sparodon durbanensis (brusher), Rhabdosargus holubi (silverbream) and Sarpa salpa (karanteen), the shad Pomatomus saltatrix and the carangid Seriola lalandi (yellowtail) (van der Elst 1988, Hutchings et al 2003). Although early papers on the subject assumed that most of these species spawned out in the Agulhas Current, as a means of moving their larvae south (Heydorn et al 1976, van der Elst 1988), more detailed studies have shown this not to be the case. For example Beckley and Connell (1998), found that the elf spawned on the shelf, in 30-50m water depth, about 5km offshore at Park Rynie (some 25-30km inside the inner edge of the Agulhas Current). See the Pomatomus sheet for more information on shad egg distribution across the shelf from the present study.      

Each species has a spawning strategy, together with early juvenile behaviour which, in combination, affords the juveniles the opportunity to recruit to their preferred nursery areas. The ability of very small juveniles of marine fishes, to move deliberately towards preferred habitat, is well established (Leis 2002). Regarding local species, nursery grounds are known to be in Cape littoral waters, for seventyfour (Ahrens, 1964), geelbek (Griffiths and Hecht 1995), karanteen (Joubert, 1981) and shad (van der Elst, 1981). A comparison of strategies in three local species, blacktail (LIIIC1), karanteen (LIIIB9) and silverbream (LIIID6), serve to illustrate the point. The present study has shown that karanteen spawn close inshore, predominantly within 500m of the shore (species sheet LIIIB9). Blacktail spawn both close inshore and on the further offshore reefs, including Aliwal Shoal (Figure 12). The spawning locality of silverbream is uncertain, since only 5 specimens were reared during this study, each as a single fish which emerged from batches of juveniles reared from eggs of blacktail and sand soldier (LIIIC5). See sheet LIIID6 for details.

Despite spawning close inshore, karanteen juveniles of 15-25mm SL have never been found in the rockpools, protected bays or estuaries of KZN. They are commonly found in bays of the eastern Cape (Joubert, 1981; Beckley, 1986). In the latter study, Beckley collected only 7 specimens, measuring 15.8-36.5mm SL. While the upper size limit might be the avoidance threshold of the net used, the lower limit is a fish in excess of 30days old (see data sheet LIIIB9). This allows that Beckley's specimens could have come from KZN spawning (note however that Strydom and d'Hotman (2005) collected juveniles as small as 8.9mm SL in the surf at Cape Padrone, at the eastern corner of Algoa Bay, indicating that spawning also occurs in eastern Cape waters). In the present study, juvenile karanteen from offshore plankton and handnet catches, have only been recognised on three occasions, twice being small specimens from the plankton net, while towing for eggs. But on one occasion, when quiet calm conditions prevailed, a loose shoal of small fish using the skiboat hull as shelter, proved to be karanteen juveniles. Details are shown in the following table:

Date

n*

Size range (SL)

Distance Offshore

Water depth (m)

9/11/2003

5/8/2004

8/8/2004

1

9 (40)

1

10.5mm

27.1-29.0mm

13.6mm

800m

4km

800m

15m

38m

15m

          * Number caught. In brackets is the estimated number in the shoal.

The shoaling behaviour seen on 5 August 2004 might suggest that congregating at the surface is a successful strategy for juveniles to move south into Eastern Cape waters. Given that inshore currents are wind dominated, and strongest at the surface, the high incidence of northeasterly winds during August to October along the KZN and Eastern Cape coasts, would contribute to south-westward transport of surface shoaling juveniles.

Adult blacktail tend to spawn a little further offshore, but inside the 30m depth contour ( LIIIC1), and their eggs were also common in the Durban Harbour mouth samples. Small juvenile blacktail abound in rocky embayments, protected reefs in the surf, and intertidal rock pools, all along the southern KZN coast in early summer (Joubert 1981). Thus, early juvenile behaviour is to make for these areas, which act as nurseries for this species.

The deliberate movement of juvenile silverbream is perhaps the most remarkable of the three. Adults are caught on deeper offshore reefs in KZN (Smith & Smith 1986), suggesting this is where they spawn, supported by five reared specimens from this study, mentioned above. But their early juveniles enter estuaries as their preferred nursery area (Wallace & van der Elst 1975). In the present study the larvae have been found to recruit to the Lovu and Mkomazi estuaries virtually all year round (Figure 7), at a constant 9.5-11mm SL, which, judging from

Figure 7
Figure 7. Recruitment pattern of Rhabdosargus holubi larvae to the Mkomazi and Lovu estuaries, from February 2005 to February 2006, sampled two-weekly on incoming spring tides.

rearing of related species such as Diplodus capensis, Rhabdosargus sarba, Sarpa salpa and Crenidens crenidens, would represent a fish of 30-35 days old. While attempting to recruit to the closed Lovu estuary, these tiny juveniles were found in the turbulent surf opposite the estuary, and even in waves running back down the sandbar’s seaward slope after failing to overtop into the estuary (Plate12). During a genuine overtopping

Plate 12

Plate 12; Netting for fish larvae in receding waves, adjacent to the closed mouth of the Lovu estuary.

event on 11 September 2002, the same net was used, and collected 5 tiny silverbream along with 8 juvenile mugilids (11-12mm SL) by netting on the estuary side of the berm, in intermittently overtopping, shallow waves (Plate 13). The deliberate presence of these estuary-recruiting species in the surf

Plate 13

Plate 13. High spring tide overtopping of waves across the berm into the Lovu estuary during a period of mouth closure.

adjacent to the estuary is demonstrated by their dominance in catches in the surf at the Lovu estuary during 2005, while the mouth was closed. About 90% of the fish larvae collected, were species that actively recruit to estuaries, and 80% were silverbream, Monodactylus sp. and mugilid juveniles combined (A Connell, unpublished). Cowley et. al. (2001) have reported similar deliberate recruitment by overtopping into closed Eastern Cape estuaries.

6.       Rainfall and spawning intensity

During September 1987, a flood event of extreme proportions occurred, during which up to 800mm of rain fell in a period of 5 days (Badenhorst et. al. 1989). The bigger rivers had flood duration periods of up to 24 hours, and divers working on an offshore pipeline, 3km off the mouth of the Mkomazi river, reported the seabed rising over 1m at the diffuser. In dives off Park Rynie, we also recorded mud lying 15cm deep in gutters in the reef at 50m depth, about 5km offshore. The nutrient load from such a deluge has never been calculated, but a striking feature of the following few years of this study was that they yielded unusually high numbers of eggs, particularly from the three most prolific pelagic egg spawners, namely the sardine, Sardinops sagax, the East coast roundherring, Etrumeus teres and the mackerel, Scomber japonicus (Connell 2001). Due to the huge variability in eggs-per-sample from this study (0 – 46235), and the enormous difference to an annual mean that a single large sample can make, the statistics are questionable. Nevertheless, a comparison of rainfall and spawning intensity, over the period of the study, does show a similar pattern (Figure 8), with an expected lag of a year, given the summer rainfall pattern and the winter/spring spawning maximum (Figure 9). Isotope studies on eggs and zooplankton are currently underway, in an effort to confirm the link.

figure 8
Figure 8: Rainfall in the Mkomazi catchment, versus mean annual egg per sample Figure 9: Mean monthly eggs per sample, averaged over 25 years.

7.       Results

7.1                  Dive log current data for the study area

Of 877 dives on which current speed and direction were logged, the number of days and percentage of 8 segments of the compass rose are shown in Figure 10. (Note that for simplicity this chart had been drawn as though the coast was aligned with True North, whereas it is in fact about 30°E of TN. Thus those currents running in the segment denoted as “north” are actually running 30°NNE, approximately parallel to the coastline, and the “south” segment is running 210°SSW). Note also that, for obvious reasons, the data is biased towards days when weather conditions favoured safe launching and diving.

7.2     Net volume and eggs per sample

From time to time, a propeller driven counter was used to calibrate the volume of water passing through the net. The results are summarised in the following table.

Number of records

Range (m3/sample)

Mean (m3/sample)

Standard Deviation

110

22.8 – 99.2

59.1

17.6

The high variability is due both to clogging of the net, variable sea conditions, and possibly, among the low values, some unseen interference with the propeller (i.e. it under-counted) during sampling.

The mean number of eggs per sample has, as mentioned in Section 6, shown a trend which appears to follow rainfall, but has essentially hinged on the successful movement into the area, of actively spawning shoals of fishes which include the sardine, east coast round herring, mackerel, the maasbanker Trachurus trachurus, a variety of Decapterus species (scads), including D. macrosoma and D. russelli, and an anchovy (BDIIIA1). The overall trend is shown in Figure 11, while the seasonal trend in spawning is shown in Figure 9 above. The months of August to October are the peak spawning months, when the three most abundant spawners, as well as most of the sparids, particularly the sandsoldier Pagellus natalensis, are spawning (see individual species sheets).

7.3     Offshore/Inshore sample analysis

As noted in the Section 2.1 of Methods, from early 1994 a second, inshore sample was collected on each launch. Comparison of the individual species distribution of eggs in the offshore and inshore samples (i.e. collected on the same day, and referred to as linked samples), gave an indication of which species spawned closer inshore (500m offshore compared with 4-5km offshore). These data are presented in a small data box on individual species sheets, with the number of samples containing at least one egg of the species, and the total number of eggs. The data from the two sample sets is not directly comparable, due to a significantly different mean number of eggs per sample, which for the offshore set, is 584, and inshore, 380 (n = 544). Reasons for this would include the obvious lack of an inshore source for the inshore sample, and higher diversity and biomass on offshore reefs. When considering individual species, and where they spawn, comparison of percentage occurrence in the two, linked sample sets provides interesting data (Table 1). Two indicator species were used to gain a relative idea of where each species spawns. The first is the kob Argyrosomus japonicus, which forms spawning aggregations on wrecks in the area, including the Produce in 30m water depth, less than 1km NW of the Aliwal Shoal (Figure 12), and the Griqualand, in 50m water depth about 15km south of Durban (Figure 2).  The other is the geelbek Atractoscion aequidens, which forms large spawning aggregations on reefs off Park Rynie. While local commercial fishermen report that they used to  be caught on reefs at 30m, they currently are mostly caught at 45-60m water depth, slightly further offshore than kob. Table 1 lists the data for all the linked samples, from the species with the highest inshore percentage at the top. Top of the list is Monodactylus falciformis, which spawns in close proximity to the surfzone, and can often be

Figure 12. Aliwal Shoal and the Produce wreck, in relation to Park Rynie and the 20m, 30m and 50m depth contours. The rectangle shows the boundaries of the Aliwal MPA.

seen feeding in loose shoals in the surfzone. The two indicator species discussed above, are highlighted in green in Table 1. In the species pages which follow, spawning is discussed in relation to the two indicator species, and the depth contours in Figure 12.

 

 

 

 

 

 

Table 1: Percentage of eggs in samples from 5km (offshore), and 0.5km (inshore), for the Park Rynie linked samples, in rank order, excluding most species with <50 eggs (1987-2011)

Code Identification number of eggs % Offshore % Inshore
FII A9 Monodactylus falciformis 2590 7 93
MII A1 TypeA Dagetichthys marginatus 68 10 90
LIIIB9 Sarpa salpa 6438 14 86
FIIIA9 Sillago chondropus 58 16 84
MIIIA4 ?Apodocreedia vanderhorsti 399 17 83
EIIIB6 Caranx sem 192 17 83
LII B6 Umbrina robinsoni 171 19 81
MIIIA8 Cynoglossus sp 4712 20 80
LIIIC1B Pachymetopon aeneum + 462 20 80
LIIID9 Rhabdosargus sarba  634 21 79
KIIIC3 Thalassoma trilobatum 327 21 79
FIIIA2A Dichistius multifasciatus 165 23 77
LIIIA6 Platycephalus indicus 105 24 76
LII A5 Aeoliscus punctulatus 125 25 75
LII B8 Scorpaenidae 49 27 73
LIIA6A Tripterodon orbis  428 27 73
LII B4 Diplodus hottentotus 333 28 72
MIIIA5 Pagusa nasuta (= Solea bleekeri) 74 28 72
LIIIC9 Pseudorhombus arsius 114 29 71
LIIIA11A Euthynnus affinis 353 32 68
LIIIA5 Cociella heemstrai 585 32 68
LIIID1 Lithognathus mormyrus 1414 33 67
EIIIB1 Upeneus guttatus 1708 34 66
LIIIE1 Parapercis robinsoni 1549 36 64
MIIIA7 Limnichthys nitidus  546 36 64
LIIIB10 Pachymetopon grande 2455 37 63
KIIIC1 Pseudanthias fasciatus 1785 40 60
FIIIA5 Chaetodon marleyi 363 40 60
EIIIB3 Pomadasys striatum 32374 40 60
LII A2 Pempheridae 801 40 60
EIIIB3A Kuhlia mugil 19 42 58
KIIIA2 Anchichoerops natalensis 161 42 58
LIIIC5 Pagellus natalensis 75730 42 58
KIIIB8 Labridae 11869 43 57
FIIIA7 Sillago cf sihama 119 43 57
LIIIF7 Unknown 304 43 57
KIIIA8 Paracaesio xanthura 1851 44 56
MIIIA2 Onigocia oligolepis & ?Thysanophrys celebica  129 45 55
CDIIIA1 Draculo celetus 236 46 54
EIIIB4 Pomadasys olivaceum 2823 46 54
MII A2 Sarda orientalis 659 47 53
CHII A1 Synodontidae 5172 47 53
FIA1 Anguilliform 79 48 52
LIIIC1  Diplodus capensis 29041 48 52
LIIIF5 Secutor insidiator 2565 49 51
MIIIA1 Rogadius portuguesus 306 49 51
LIIIF2 Chaetodon spp. 3855 49 51
BFIIIA1 Stolephorus holodon 268 49 51
G II A1 Hilsa kelee 52 50 50
EIIIB8 Gerres acinaces 280 51 49
LIIIG7 Crossorhombus valderostratus 15325 51 49
KIIIB2 Cyprinocirrhites polyactis 377 51 49
FIIIA2 Dinoperca petersi 269 54 46
KIIIA4 Serranus knysnaensis 599 55 45
EIIIA6 Decapterus spp &Trachurus trachurus 53082 56 44
LIIIE7 Acanthurus mata 9134 56 44
KIIIA6 Brotula multibarbata 1243 56 44
CMI A2 Zebrais regani 75 56 44
LIIIB7 Myxus capensis + other mugilids 397 56 44
LIIIB4  Epinephelus spp 46 57 43
FII A1 Sardinops sagax 153669 57 43
KIIIA10 Parupeneus spp. 14694 57 43
KIIIA1 Chirodactylus jessicalenorum  4573 57 43
KIIIB2A Cirrhitichthys oxycephalus 751 58 42
FIIA7 Oplegnathus conwayi 75 59 41
LII A3 Aulostomus chinensis 200 59 41
FIIIA3 Paralichthodes algoensis 94 60 40
LIIIA11 Auxis rochii 18905 60 40
K III B3A Gymnocranius cf griseus 390 60 40
LIIB7 Apistus carinatus 92 61 39
CMI A1 Ostraciidae 593 61 39
KIIIB3 Priacanthus hamrur 1086 61 39
MII A5 Pempheridae 370 61 39
FIIIA4  Pomatomus saltatrix 3810 62 38
DIIIA3 Callionymidae 991 63 37
FII B1 Neoscorpis lithophilus 152 63 37
DI A1 Anguilliform 515 64 36
HII A3 ?Choridactylus natalensis 31 65 35
LIIIF9  Bothidae 274 65 35
LIIID2 Epinephelus spp. 784 67 33
HII A4 Suarida undusquamis 1183 67 33
EIIIA2 Oplegnathus robinsoni 88 68 32
FIIIA6 Pomacanthus rhomboides 260 68 32
DII A1 Etrumeus teres 65917 69 31
LIIIG4 Acanthurus triostegus 4212 69 31
EIIA2 Seriola dumerili & S. lalandii 99 70 30
LII B5 Lepidotrigla faurei 1324 70 30
KIIIB7 Calotomus carolinus & Labridae 1068 71 29
LIIIE9 Pseudorhombus elevatus 241 71 29
BKIIIA2 Scarus spp 5464 71 29
HI A3 Fistularia spp 197 73 27
LIIIE4 Antigonia rubescens 243 73 27
LIIIA8  Argyrosomus japonicus 4311 73 27
LIIIF3 Cubiceps pauciradiatus 159 74 26
EII A3  Centroberyx spinosus 1737 74 26
LII A7 Scomber japonicus 108855 75 25
LIA4 Brama brama 76 75 25
BDIIIA1 Engraulus encrasicolus & Encrasicholina punctifer 19317 76 24
LIIIA4 Umbrina canariensis + 560 76 24
GIA1 Anguilliform 85 76 24
EIIIB6A Myripristis berndti 250 80 20
FII A4  Coryphaena hippurus 307 81 19
EIIIA9 Emmelichthys struhsakeri 941 81 19
CDI A2 Scomberesox saurus 119 82 18
LII B3 Aulacocephalus temmincki  324 83 17
LI A1 Caristius spp. 126 83 17
KIIIB1 Pristipomoides sieboldii & Paracaesio sordida 6492 84 16
KIIIA9 Paracaesio xanthura & Caesio cf caerulaurea 60050 84 16
LII A6 Atractoscion aequidens 3040 85 15
DIIIA1 Vinciguerria nimbaria 510 85 15
ABHIIIA1 Parascorpaena mossambica + 1641 85 15
EIIIA8 Centroberyx druzhinini 91 86 14
EIIIA1 Sphyraena jello 153 88 12
BHIIA1 Unknown 109 88 12
BKIIIA1 Carapidae 1396 88 12
LIIIE8 Heniochus acuminatus 350 89 11
FII A5 Centrolophus niger 1227 89 11
FIIIA1 Acanthistius joanae 793 90 10
EII A4  Seriola dumerili & S. rivoliana 640 90 10
EIIIB5 Pristigenys niphonia 127 92 8
LIIIA11B Katsuwonus pelamis 153 92 8
FII A8 Plagiogeneion rubigonosum 53 92 8
LIIIE10 Zebrasoma gemmatum 177 93 7
DIIIA4 Unknown 1472 95 5
KIIIB5 Unknown 79 97 3
BLIIIA2 Scorpaenidae 649 100 0

 

7.4     The most common eggs from Durban Bay and offshore

The following two tables summarise the most common eggs, based on total numbers found over the duration of each study.

Table 2: The most common eggs collected in the mouth of Durban Harbour on outgoing spring tides, from May 1990 to December 1994, arranged in order of abundance (number of samples =77).

n

Code

Species or description

n

Code

Species or description

10531

LIIIE3A

Acanthopagrus vagus

13

DIIIA3

Callionymus marleyi

9263

GII A1

Hilsa kelee

11

FIIIA2A

Dichistius multifasciatus

5484

EIIIB2

Pomadasys commersonnii

11

LIIIA5

Cociella heemstrai

4727

LIIID9

Rhabdosargus sarba

11

LIIIC9

Pseudorhombus arsius

4333

LIIIA7

Liza dumerilii and other mugilids

10

LII A2

Pempheris schwenkii

3646

DI A1

Anguilliformes

8

KIIIB7

Labridae

1799

KIIIB2A

Cirrhitichthys oxycephalus

8

LIIIF7

Unknown

1312

LIIIC1

Diplodus capensis, blacktail

7

LIIIE1A

Parapercis sp

1142

LIIIF5

Secutor ruconius

7

LIIIE3

Sciaenidae

986

BFIIIA1

Stolephorus holodon

7

LII A3

Aulostomus chinensis

948

KIIIB9

Ambassis dussumieri

6

MII A1

Dagetichthys marginatus

874

EIIIB8

Gerres acinaces

6

LII B5

Lepidotrigla faurei

853

DII A2

Thryssa setirostris

5

EIIIA6

Trachurus trachurus & Decapterus spp.

637

LIIIE11

Crenidens crenidens

5

KIIIA1

Chirodactylus jessicalenorum

306

MIIIA2

Thysanophrys celebica

5

LIIIA11

Auxis rochei

249

LIIIG4

Acanthurus triostegus

5

LIIIC3

Pseudorhombus sp.

130

KIIIB8

Halichoeres lapillus +

4

BDIIIA1

Engraulis encrasicolus & Encrasicholina punctifer

123

LIIID2

Epinephelus spp.

4

KIIIA6

Brotula multibarbata

118

LIIIF2

Chaetodon dolosus & C. blackburni

4

LII B9

Liza tricuspidens

114

LIIIG7

Arnoglossus

4

MIIIA5

Solea turbynei (bleekeri)

94

LIIIE1

Parapercis robinsoni

3

BKIIIA2

Scarus rubroviolaceus

66

MIIIA8

Cynoglossus sp

3

EIIIB4

Pomadasys olivaceum

65

KIIIC1

Pseudanthias squamipinnis

3

FIIIA4

Pomatomus saltatrix

57

KIIIA10

Parupeneus spp.

3

HI A2

Fistularia petimba

56

KIIIA2

Anchichoerops natalensis

3

HII A1

Tetraodontidae

54

DII A1

Etrumeus teres

3

KIIIA9

Caesio cf caerulaurea

53

EIIIB1

Upeneus guttatus

2

FI A1

 Anguilliformes

50

HII A4

Saurida undusquamis

2

HII A3

Choridactylus natalensis

39

FII A9

Monodactylus falciformis

2

KIIIC3

Thalassoma trilobatum

38

FIIIA4B

Sillago cf sihama

2

LI A2

Trichiurus lepturus

34

LII A5

Aeoliscus punctulatus

2

LIIIA4

Umbrina canariensis (?+ others)

30

LIIIA6

Platycephalus indicus

2

MIIIA7

Limnichthys nitidus

28

LIIIB10

Pachymetapon grande

1

CMI A1

Tetrosomus concatenates

25

MII A2A

Asseragodes heemstrai

1

KIIIB3

Priacanthus hamrur

18

FII A1

Sardinops sagax

1

FIIIA2

Dinoperca petersi

17

DIIIA1

Phosichthyidae

1

LII A7

Scomber japonicus

17

KIIIB9A

Unknown

1

LII B6

Umbrina robinsoni

16

CHII A1

Trachinocephalus myops

1

LII B8

Scorpaenidae

16

LIIIB9

Sarpa salpa

1

LIIIE7

Acanthurus mata

15

LIIIC5

Pagellus natalensis

1

MI A1

Lampridiformes

14

KIIIA4

Serranus knysnaensis

1

MIIIA1

Rogadius portuguesus

14

KIIIA8

Paracaesio xanthura

1

MIIIA4

?Apodocreedia vanderhorsti

14

LII B4

Diplodus hottentotus

     

Table 3: The top 154 most common eggs from Park Rynie samples, collected from 1987 to 2011 inclusive, arranged in order of abundance.

n

Code

Species or description

n

Code

Species or description

234342

FII A1

Sardinops sagax

355

LIIIE8

Heniochus acuminatus

194601

LII A7

Scomber japonicus

346

MIIIA1

Grammoplites portuguesus

114385

DII A1

Etrumeus teres & E. whiteheadi

341

FIIA4

Coryphaena hippurus

95583

LIIIC5

Pagellus natalensis

341

LIIB4

Diplodus hottentotus

61038

KIIIA9

Paracaesio xanthura & Caesio caerulaurea

326

LIIIF7

Unknown

56310

EIIA6

Trachurus & Decapterus

298

KIIIB1A

Paracaesio sordida

43420

EIIIB3

Pomadasys striatum

294

LIIIE9

Pseudorhombus elevatus

30777

LIIIC1

Diplodus capensis

291

LIIIF9

Engyprosopon grandisquama

24988

BDIIIA1

Engraulis encrasicolus & Encrasicolina punctifer

274

FIIIA6

Pomacanthus rhomboides

21688

LIIIA11

Auxis rochii

250

EIIIB6A

Myripristis berndti

21593

LIIIG7

Crossorhombus valdirostratus

244

FIIIA2

Dinoperca petersi

16853

KIIIA10

Parupeneus fraserorum & P. rubescens

237

CDIIIA1

Draculo celetus +

12443

KIIIB8

Halichoeres lapillus +

232

HIA3

Fistularia commersonii & F. petimba

9382

LIIIE7

Acanthurus mata

229

MIIA1

Dagetichthys marginatus

8237

LIIIA8

Argyrosomus japonicus

223

EIIIB6

Caranx sem

7947

LIIIA5

Secutor insidiator

216

MIIA6

Cynoglossus lida

7179

LIIIB9

Sarpa salpa

211

LIIIE5

Macroramphosus scolopax

6699

KIIIB1

Paracaesio sordida & Pristipomoides sieboldii

208

LIIA3

Aulostomus chinensis

6652

BKIIIA2

 Scarus rubroviolaceus & S. ghobban

205

CDIA2

Scomberesox saurus

6590

MIIIA8

Cynoglossidae

204

LIIB6

Umbrina robinsoni

6235

CHII A1

Trachinocephalus myops

193

FIIIA2A

Dichistius multifasciatus

5933

FIIIA4

Pomatomus saltatrix

183

EIIIB5

Pristigenys niphonia

4933

KIIIA1

Chirodactylus jessicalenorum

181

KIIIA2

Anchichoerops natalensis

4929

EIIIB4

Pomadasys olivaceum

181

MIIIA2

Onigocia oligolepis +

4647

KIIIA8

Paracaesio xanthura

179

LIIIE10

Zebrasoma gemmatum

4483

LIIIG4

Acanthurus triostegus

173

LIIA11B

Katsuwonus pelamis

4268

LIIIF2

Chaetodon spp.

169

LIIIF3

Cubiceps pauciradiatus

3386

FII A9

Monodactylus falciformis

163

LIIIC9

Pseudorhombus arseus

3213

LIIA6

Atractoscion aequidens

160

FII B1

Neoscorpis lithophilus

2830

ABHIIIA1

Scorpaenopsis possi & Parascorpaena mossambica

157

EIIIA1

Sphyraena jello

2810

LIIIB10

Pachymetopon grandis

145

MIIIA5

Pagusa nasuta

2330

KIIIC1

Pseudanthias fasciatus & P. squamipinnis

143

FIA1

Anguilliform

2112

LIIB5

Lepidotrigla faurei

138

FIIIA7

Sillago cf sihama

2095

EII A3

Centroberyx spinosus

134

LIA1

Caristius sp

2062

EIIIB1

Upeneus guttatus

131

LIIIA6

Platycephalus indicus

1853

HIIA4

Saurida undosquamis

130

LIIA5

Aeoliscus punctulatus

1812

LIIIE1

Parapercis robinsoni +

127

LIIB9

Myxus capensis & Liza tricuspidens

1704

DIIIA4

Callionymidae

123

GIA1

Anguilliform

1586

KIIIB7

Labridae & Calotomus carolinus

119

BHIIA1

Unknown

1527

BKIIIA1

Carapidae

118

EIIIA8

Centroberyx druzhinini

1517

DIIIA3

Callionymus marleyi

117

EIIIA2

Oplegnathus robinsoni

1452

LIIID1

Lithognathus mormyrus

110

FIIIA2

Paralichthodes algoensis

1426

KIIIA6 Brotula multibarbata

109

EIIA2

Seriola dumerili & S. lalandi

1364

FIIA5

Centrolophus niger

108

LIIB7

Apistus carinatus

1230

KIIIB3

Priacanthus hamrur

107

CMIA2

Zebrias cf regani

1220

LIIID9

Rhabdosargus sarba

104

KIIIB5

Unknown

1217

FIIIA1

Acanthistius joanae

89

LIIIB4

Epinephelus malabaricus & E. andersoni

1125

LIIIA4

Umbrina canariensis

88

FIIA7

Oplegnathus conwayi

1019

EIIIA9

Emmelichthys struhsakeri

84

LIA4

Brama brama

1008

BFIIIA1

Stolephorus holodon

83

LIA2

Trichiurus lepturus

953

MII A2

Sarda orientalis

81

FIIA8

Plagiogeneion rubiginosum

931

LIIA2

Pempheris schwenkii

73

HIIA3

Choridactylus natalensis

890

KIIIB2A

Cirrhitichthys oxycephalus

68

CHIA1

Oxyporhamphus micropterus

825

LIIID2

Epinephelus rivulatus +

68

CLIIA2

Myctophiform

786

CMIA1

Ostraciidae

64

CDIA1

Regalecus glesne

783

KIIIA4

Serranus knysnaensis

63

GIIA1

Hilsa kelee

775

DIIIA1

Vinciguerria nimbaria

60

FIIIA9

Sillago chondropus

748

LIIIA5

Cociella heemstrai

59

LIIB8

? Scorpaenidae

689

EIIA4

Seriola spp

48

MIA3

Diodon holocanthus

663

MIIIA7

Limnichthys nitidus

46

LIIIE3A

Acanthopagrus vagus

652

BLIIIA2

 Scorpaenidae

43

EII A1

Pentaceros capensis

634

DIA1

Anguilliforms

42

EIII A3A

Sphyraena sp

570

LIIIB7

Liza dumerilii +

42

HIIA2

Scorpaenidae

523

LIIIA11A

Euthynnus affinis

40

ABKIIIA1

Carapidae

498

LIIA6A

Tripterodon orbis

39

FIIA3

Hyperoglyphe antarctica

466

KIIIB2

Cyprinocirrhites polyactis

36

LIIIG1

Bothidae

465

LIIICiB

Pachymetopon aeneum

35

KIIIC2

Anthiinae (sea goldie)

456

EIIIB4A

Megalaspis cordyla

33

LIIA4A

Scomberomorus plurilineatus

443

MIIA5

Parapriacanthus ransonneti

33

LIIB1

Champsodon capensis

443

MIIIA4

?Apodocreedia vanderhorsti

32

LIIIB3

Sparidae

427

EIIIA3

Carangidae

31

BLIIIA1

Helicolenus dactylopterus

425

FIIIA5

Chartodon marleyi

30

ABHIIIA2

Scorpaenidae

413

EIIIB8

Gerres acinaces

30

LIIIC3

Pseudorhombus sp.

408

LIIB3

Aulacocephalus temminckii

27

LIIID4

Cephalopholis sonnerati

380

KIIIC3

Thalassoma trilobatum

26

LIIIE3

?Argyrosomus thorpei

365

KIIIB3A

Gymnocranius griseus

25

EIIIA1A

Kyphosus bigibbus

359

LIIE4

Antigonia rubescens

25

LIIIC4

Malacanthus brevirostris

 

8.       Voucher specimens.

Representative samples of all reared larvae and juveniles have been archived, and are deposited in the SAIAB collection at Rhodes University in Grahamstown.

9.       Acknowledgements

Professor Paul Hebert and his team, particularly Tyler Zemlak and Dirk Steinke, at the University of Guelph, Canada, whose assistance with barcoding of larvae has been a mainstay to identifying larvae, thereby contributing hugely to the value of the dataset.

The MODIS Project and the Active Archive Centre at the Goddard Space Flight Centre, Greenbelt, MD 20771, for the production of MODIS data, and Bertrand Saulquin for providing these data (Plates 3, 9 and 10).

Park Rynie Skiboat Club, for use of their launch facilities at Rocky Bay.

The CSIR, which funded the sample collection from 1987 to 2003, and for permission to use Plates 4-8 in the Introductory Notes.

The Oceanographic Research Institute, Durban, for allowing me to collect eggs from the main tank, and for collaborative work with staff, particularly Dr Pat Garratt.

The South African Institute for Aquatic Biodiversity, Grahamstown, for ichthyological expertise, and for becoming involved in the "Barcode of fishes" initiative.

Mike, Valda and Alan Fraser, for their interest and insights, fish DNA samples, and for assisting with sample collection while I was incapacitated between March and November 2006.

My network of fish collectors, for DNA barcoding of adult fishes, including Knud Sorensen, captain of the trawler Ocean Spray, Tim McClurg and Steven Weerts at CSIR Durban, Rob Broker and Paul Buchel at KZN Wildlife, Sheldon Dudley at Natal Sharks Board, Simon Chater, Sean Fennessy and Bruce Mann at ORI, Andre Bok at I&J in Danger Point, Ken and Larry Hutchings at University of Cape Town and, especially, Rob Cooper at Marine & Coastal Management in Cape Town.

Dr Phil Heemstra and Elaine Heemstra, for ichthyological expertise and many DNA samples.

FC (Billy) Clark, whose experience of the sea and fishing commercially for "74" and other species of linefish in the 1950’s and 60’s, aboard his vessel The Plough, along the KZN and eastern Cape coast, has added interesting anecdotal information to some of the species sheets.

10.            Literature

Ahlstrom EH, Butler JL & Sumida BY. 1976. Pelagic stromateoid fishes (Pisces, Perciformes) of the eastern Pacific: kinds, distributions, and early life histories and observations of five of those from the Northwest Atlantic. Bulletin of Marine Science 26: 285-402.

Ahlstrom EH & Counts RC. 1958. Development and distribution of Vincihuerria lucetia and related species in the eastern Pacific. United States Fisheries Bulletin 58: 363-416.

Ahrens R L 1964. A preliminary report on the biology of the seventy- four Polysteganus undulosus (Regan) in Natal waters. Unpublished M.Sc. thesis. University of Natal, Durban: pp 1-77.

Anders AS. 1975. Pilchard and anchovy spawning along the Cape east coast. South African Shipping News and Fishing Industry Review 30: 53-57.

Badenhorst P, Cooper JAG, Crowther J, Gonsalves J, Grobler NA, Illenberger WK, Laubscher WI, Mason TR, Moller JP, Perry JE, Reddering JSV and van der Merwe L. 1989. Survey of September 1987 Natal floods. South African National Scientific Programmes Report 164: 1-135.

Beckley LE. 1986. The ichthyoplankton assemblage of the Algoa Bay nearshore region in relation to coastal zone utilisation by juvenile fish. South African Journal of Zoology 21: 244-252.

Beckley LE and Connell AD. 1996. Early life history of Pomatomus saltatrix off the East coast of South Africa. Marine and Freshwater Research 47: 319-322.

Beckley LE and Hewitson JD. 1994. Distribution and abundance of clupeoid larvae along the east coast of South Africa in 1990/1991. South African Journal of Marine Science 14: 205-212.

Beckley LE and Naidoo AK. 2003. Exploratory trials with light-traps to investigate settlement stage fishes in subtropical, coastal waters off South Africa. South African Journal of Zoology 38: 333-342.

Begg G. 1978. The estuaries of Natal. Natal Town and Regional Planning, 41: 1-657.

Blaber SJ, Cyrus DP and Whitfield AK. 1981. The influence of zooplankton food resources on the morphology of the estuarine clupeid Gilchristella aestuarius (Gilchrist 1914). Environmental Biology of Fishes 6: 351-355.

Brownell CL. 1979. Stages in the early development of 40 marine fishes with pelagic eggs from the Cape of Good Hope. Ichthyological Bulletin of the J.L.B. Smith Institute of Ichthyology. 40: 84p.

Carpenter KE. 2001. Lethrinidae. In: Carpenter KE and Niem VH.(Eds.) FAO species identification guide for fisheries purposes. The living marine resources of the Western Central Pacific. Volume 5. Bony fishes Part 3(Menidae to Pomacanthidae). Rome FOA. pp. 3004-3050.

Carter RA. 1977. The distribution of calanoid copepods in the Agulhas Current system off Natal, South Africa. MSc Thesis, University of Natal, 165pp.

Carter R and D’Aubrey J. 1988. Inorganic nutrients in Natal continental shelf waters. In: Coastal Ocean Studies off Natal, South Africa. E.H. Schumann (ed.) Berlin. Springer Verlag, pp 132-151.

Carter R and Schleyer M H. 1988. Plankton distributions in Natal coastal waters. In: Coastal Ocean Studies off Natal, South Africa. E.H. Schumann (ed.) Berlin. Springer Verlag, pp 152-177.

Cohen DM 1986. Family No 88 Gadidae; Family No 89: Merlucciidae; Family No 90: Moridae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes:324-328.

Collette BB. 1986. Family No. 249: Scombridae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 831-838.

Connell AD. 1985. Neuston netting. In: Environmental studies at Richards Bay prior to the discharge of submarine outfalls: 1974-1984. CSIR Durban Marine Research Group Report. Pp.175-177.

Connell AD. 1989. Neuston netting. In: Environmental studies in the Richards Bay Offshore Outfalls Region. Part 2. Surveys made between 1985 and 1988. CSIR Report EMA-C 8984 pp141-144.

Connell AD. 1996. Sea fishes spawning pelagic eggs in the St Lucia estuary. South African Journal of Zoology 31:37-41.

Connell AD 1997. Seasonal trends in sardine spawning at Park Rynie, Kwazulu-Natal south coast. WOSAS – Workshop on Southern African sardine, (eds.) M. Barange & C. van der Lingen. Sea Fisheries Research Institute Benguela Ecology Programme Report 29:29-33.

Connell AD. 1997a. Neuston netting. In: Environmental studies in the Richards Bay Offshore Outfalls Region. Report No. 10. Surveys made during 1996. CSIR Report ECP-97-011, Part 7: 147-154.

Connell AD. 1998. Neuston netting. In: Environmental studies in the Richards Bay Offshore Outfalls Region. Report No. 11. Surveys made during 1997. CSIR Report ENV/ECP/EXT:97-JEC17, Part 7: 147-154.

Connell AD. 2001. Pelagic eggs of marine fishes from Park Rynie KwaZulu-Natal, South Africa: seasonal spawning patterns of the three most common species. African Zoology 36: 197-204.

Connell AD. 2003. Neuston netting. In: Environmental studies in the Richards Bay Offshore Outfalls Region. Report No. 16: Surveys made during 2002. CSIR Report ENV-D-C-2003-003, Part 7: 153-158.

Connell AD. 2010. A 21-year ichthyoplankton collection confirms sardine spawning in KwaZulu-Natal waters., African Journal of Marine Science 32: 331-336.

Connell AD, Heemstra PC and Garratt PA. 1999. Eggs and larvae of the santer Cheimerius nufar from KwaZulu-Natal. South African Journal of Marine Science 21: 41-50.

Cowley PD, Whitfield AK and Bell KNI. 2001. The surfzone ichthyoplankton adjacent to an intermittently open estuary, with evidence of recruitment during marine overwash events. Estuarine Coastal and Shelf Science 52: 339-348.

Cyrus DP, Vivier L and MacKay CF. 2005. Lake St Lucia fish survey- December 2004. Coastal Research Unit of Zululand – Investigational Report No. 118: 1-27.

Delsman HC. 1922. Fish eggs and larvae from the Java Sea. 2. Chirocentrus dorab. Treubia 3: 38-46.

Delsman HC. 1931. Fish eggs and larvae from the Java Sea. 17. The genus Stolephorus. Treubia, 13: 217-243.

Eschmeyer WN. 1986. Family 149: Scorpaenidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: 463-478.

Fahay MP. 1983. Guide to the early life stages of marine fishes occurring in the western North Atlantic Ocean, Cape Hatteras to the southern Scotian Shelf. Journal of the North West Atlantic Fisheries Science 4: 423 p.

Fahay MP & Markle DF 1983. Gadiformes: Development and Relationships. Pp 265-283. In: American Society of Ichthyologists and Herpetologists Special Publ. 1. Allen Press 760 pp.

Flemming B and Hay R. 1988. Sediment distribution and dynamics on the Natal continental shelf. In: Coastal Ocean Studies off Natal, South Africa. E.H. Schumann (ed.) Berlin. Springer Verlag, pp 47-80.

Fricke R 1986. Family No. 239: Callionymidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 770-774.

Garratt, PA. 1985. The Offshore Linefishery of Natal: II: Reproductive biology of the sparids Chrysoblephus puniceus and Cheimerius nufar. Investigational Report of the Oceanographic Research Institute of South Africa 63: 21 pp.

Garratt, PA. 1991. Spawning behaviour of Cheimerius nufar in captivity. Environmental Biology of Fishes 31: 345-353.

Garratt PA. 1993. Spawning of the river bream, Acanthopagrus berda, in Kosi estuary. South African Journal of Zoology 28: 26-31.

Gilchrist JDF and Thompson WWF 1914. Descriptions of fishes from the coast of Natal (Part IV). Annals of the South African Museum 13: 65-95.

Griffiths MH and Hecht T. 1995. On the life-history of Atractoscion aequidens, a migratory sciaenid off the east coast of southern Africa. Journal of Fish Biology 47: 962-985.

Haedrich RL 1986. Family No. 254: Stromateidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 842-846.

Hardy JD. 2006. Chiasmodontidae: Swallowers. In: Richards WJ (Ed.) Early life stages of Atlantic Fishes: An Identification Guide for the Western Central North Atlantic. Chapter 162: 1901-1915.

Harris SA and Cyrus DP. 1998. Composition, abundance and seasonality of fish larvae in the mouth of Durban Harbour, Kwa-Zulu Natal, South Africa. South African Journal of Marine Science 21: 19-39

Harrison TH 2003. Biogeography and community structure of fishes in South African estuaries. PhD Thesis, Rhodes University Grahamstown, South Africa. 229 pp.

Hebert PDN, Ratnasingham S and deWaard JR 2003. Barcoding animal life: Cytochrome C oxidase subunit 1 divergencies among closely related species. Proc. R. Soc London Ser. B 270: S596-S599.

Hecht J. 2003. Every species has a ‘barcode’. New Scientist 22March 2003:pg 14.

Heemstra PC 1986. Family No. 234: Mugiloididae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 739-741.

Heemstra PC 1986a. Family No. 229: Champsodontidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 734-735.

Heemstra PC 1986b. Family No. 261: Cynoglossidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 865-868.

Heemstra PC 1986c. Family No. 230: Uranoscopidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 735-736.

Heemstra PC 1986d. Family No. 126: Berycidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 409-410.

Heemstra PC 1986e. Family No. 79: Synodontidae (revised). In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 270-273.

Heemstra PC 1986f. Family No. 208: Caristiidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 636-637.

Heemstra PC 1986g. Family No. 206 Oplegnathidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 632-633.

Heemstra PC 1986h. Family No. 122 Regalecidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: p. 403.

Heemstra PC and Gon O. 1986. Family No. 262: Soleidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 868-874.

Heemstra PC and Heemstra E. 2004. Coastal Fishes of South Africa. SAIAB and NISC, Grahamstown.

Heemstra PC and Randell JE. 1986. Family No. 165: Serranidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 509-537.

Henseley DA. 1986. Family No. 259: Bothidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 854-863.

Heydorn AEF, Bang ND, Pearce AF, Fleming BW, Carter RA, Schleyer MH, Hughes GR, Bass AJ, van der Elst RP. Crawford RJM and Shelton PA 1978. Ecology of the Agulhas current region: an assessment of biological responses to environmental parameters in the south-west Indian Ocean. Transactions of the Royal Society of South Africa 43: 151-190.

Horn MH. 1984. Stromateoidei: development and relarionships pp 620-628. In: American Society of Ichthyologists and Herpetologists Special Publication 1.

Houde ED, Almatar S, Leak JC and Dowd CE. 1986. Ichthyoplankton abundance and diversity in the western Arabian Gulf. Kuwait Bulletin of Marine Science 8: 107-393.

Hunter IT. 1988. Climate and Weather off Natal. In: Coastal Ocean Studies off Natal, South Africa. E.H. Schumann (ed.) Berlin. Springer Verlag, pp 81-100.

Hutchings L, Beckley LE, Griffiths MH, Roberts MJ, Sundby S and van der Lingen C. 2003. Spawning on the edge: spawning grounds and nursery areas around the southern African coastline. Marine and Freshwater Research 53: 307-318.

Ikeda T and Mito S. 1988. Pelagic fish eggs pp 999-1083. In: Okiyama (ed) 1988.

Johnson RK and Keene MJ. 1986. Family No. 228: Chiasmodontidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 731-734.

Jones DL, Lara MR and Richards WJ. 2006. Labridae: Wrasses. In: Richards WJ (Ed.) Early life stages of Atlantic Fishes: An Identification Guide for the Western Central North Atlantic. Chapter 157: 1835-1871.

Joubert CSW. 1981. Aspects of the biology of five species of inshore reef fishes on the Natal coast, South Africa. Investigational Report. Oceanographic Research Institute 51: 1-16.

Knapp LW. 1986. Family No. 155: Platycephalidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 482-486.

Lamkin JT. 2006. Nomeidae: Driftfishes. In: Richards WJ (Ed.) Early life stages of Atlantic Fishes: An Identification Guide for the Western Central North Atlantic. Chapter 197: 2255-2271.

Leis JM. 1977. Development of the eggs and larvae of the slender mola, Ranzania laevis (Pisces, Molidae). Bulletin of Marine Science 27: 448-466.

Leis JM, Carson-Ewart BM and Cato DH. 2002. Sound detection in situ by the larvae of a coral reef damselfish. Marine Ecology Progress Series 232: 259-268.

Leis JM, and Carson-Ewart BM. 2003. Orientation of pelagic larvae of coral reef fishes in the ocean. Marine Ecology Progress Series 252: 239-253.

Leis JM and Rennis DS. 1983. The larvae of Indo-Pacific coral reef fishes. New South Wales University Press Sydney. 269pp.

Leis JM and Trnski T. 1989. The larvae of Indo-Pacific shorefishes. New South Wales University Press Sydney. 371pp.

Leis JM, Trnski T and Beckley 2002. Larval development of Pagellus natalensis and what larval morphology indicates about relationships in the perciform fish family Sparidae. Mar. Freshwater Res. 53: 367-376.

Lutjeharms JRE and Connell AD. 2000. The Natal Pulse and inshore counter-currents off the South African east coast. South African Journal of Science 85: 533-535.

Lutjeharms JRE, Cooper J and Roberts M. 2000. Upwelling at the inshore edge of the Agulhas Current. Continental Shelf Research 20: 737-761.

Lutjeharms JRE and Roberts HR. 1988. The Natal Pulse: an extreme transient of the Agulhas Current. Journal of Geophysical Research 93 (C1): 631-645.

Lyczkowski-Shulz J. 2006. Molidae: Molas, ocean sunfishes. In: Richards WJ (Ed.) Early Stages of Atlantic Fishes: An Identification Guide for the Western Central North Atlantic. Chapter 215:2457-2465.

Mann BQ. 2007. Reassessment of the seventy-four Polysteganus undulosus stock after a 10-year moratorium. Oceanographic Research Institute, Durban, ORI Unpublished Report 244: pp. 1-22.

Marcus LR. 2006. Caristiidae: Manefish. In: Richards WJ (Ed.) Early Stages of Atlantic Fishes: An Identification Guide for the Western Central North Atlantic. Chapter 135:1539-1543.

Matarese AC, Kendell AW, Blood DM and Vinter BM. 1989. Laboratory guide to early life history stages of Northeast Pacific fishes. NOAA Technical Report NMFS 80: 649p.

McKay RJ. 1986. Family No. 198: Sillaginidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 615-616.

Merrett NR. 2006. Bathygadidae & Macrouridae. In: Richards WJ (Ed.) Early Stages of Atlantic Fishes: An Identification Guide for the Western Central North Atlantic. Chapter 43:595-616.

Mito S 1961. Studies on the pelagic fish eggs and hatched larvae found in the adjacent waters of Japan. Records of Oceanographic Works in Japan. Special No. 5. Pp 71-94.

Mito S 1961a. Pelagic fish eggs from Japanese waters – I. Clupeina, Chanina, Stomiatina, Myctophida, Anguillida, Belonida and Syngnathida. Scientific Bulletin, Faculty of Agriculture, Kyushu University 18(3):285-310 plus plates 20-34 (in Japanese, English summary).

Mito S 1961b. Pelagic fish eggs from Japanese waters – II. Lamprida, Zeida, Mugilina, Scombrina, Carangina and Stromateina. Scientific Bulletin, Faculty of Agriculture, Kyushu University 18(4):451-466 plus plates 39-43 (in Japanese, English summary).

Mito S 1962. Pelagic fish eggs from Japanese waters – V. Callionymina and  Ophidiina. Scientific Bulletin, Faculty of Agriculture, Kyushu University 19(3):377-380 plus plates 11-12 (in Japanese, English summary).

Mito S 1963. Pelagic fish eggs from Japanese waters – III. Percidae Japanese Journal of Ichthyology 11(1-2):39-64 plus plates 1-18 (in Japanese, English summary).

Mito S 1963b. Pelagic fish eggs from Japanese waters – IX. Echeneida and Pleuronectida. Japanese Journal of Ichthyology 11(3-6):81-102 plus plates 29-41 (in Japanese, English summary).

Mito S 1963c. Pelagic fish eggs from Japanese waters –X. Gadida and Lophiida. Japanese Journal of Ichthyology 11(3-6):103-113 plus plates 42-45 (in Japanese, English summary).

Moser HG, Richards WJ, Cohen DM, Fahay MP, Kendall AW and Richardson SL. 1984. Ontogeny and systematics of fishes. Special Publication 1, American Society of Ichthyology and Herpetology. Allen Press 760p.

Nakamura I. 1986. Family No. 248: Trichiuridae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 829-830.

Okiyama M. (ed) 1988. An atlas of the early stage fishes in Japan. Tokai University Press, Tokyo. 1154pp. (in Japanese).

Olney JE. 2006. Lophotidae: Crestfish & unicornfish. In: Richards WJ (Ed.) Early life stages of Atlantic Fishes: An Identification Guide for the Western Central North Atlantic. Chapter 82: 999-1003.

Olney JE and Richards WJ. 2006. Trachipteridae: Dealfishes and Ribbonfishes. In: Richards WJ (Ed.) Early life stages of Atlantic Fishes: An Identification Guide for the Western Central North Atlantic. Chapter 86: 1019-1026.

Pearce AF. 1977. The shelf circulation off the east coast of South Africa. CSIR Research Report 361: 1-220.

Pearce AF. 1978. Seasonal variations of temperature and salinity on the northern Natal continental shelf. South African Geographical Journal 60: 135-143.

Pearce AF, Schumann EH and Lundie GSH. 1979. Features of the shelf circulation off the Natal coast. South African Journal of Science 74: 328-331.

Preston-Whyte RA. 1969. Sea breeze studies in Natal. South African Geographical Journal 51: 38-49.

Randall JE. 1961. A contribution to the biology of the convict surgeonfish of the Hawaiian Islands, Acanthurus triostegus sandvicensis. Pacific Science 15: 215-272.

Randall JE. 1986. Family No. 220: Labridae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 683-706.

Richards WJ. 2006. Scombridae: Mackerels and Tuna. In: Richards WJ (Ed.) Early life stages of Atlantic Fishes: An Identification Guide for the Western Central North Atlantic. Chapter 191: 2187-2227.

Roberts MJ, Zemlak T and Connell AD. 2011. Cyclonic eddies reveal Oegopsida squid egg balloon masses in the Agulhas Current, South Africa. African Journal of Marine Science 33: 239-246.

Schumann EH. 1988. Physical Oceanography off Natal. In: Coastal Ocean Studies off Natal, South Africa. E.H. Schumann (ed.) Berlin. Springer Verlag, pp 101-130.

Senou H. 2001. Sphyraenidae. In: Carpenter KE and Niem VH.(Eds.) FAO species identification guide for fisheries purposes. The living marine resources of the Western Central Pacific. Volume 6. Bony fishes Part 3(Labridae to Latimeriidae). Rome FOA. pp. 3685-3697.

Shao KT., Yang RS., Chen KC and Lee YS. 2001. An identification guide of marine fish eggs from Taiwan. Institute of Zoology Academia Sinica 176p.

Smith MM. 1986. Family No. 207: Bramidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 633-636.

Smith MM. 1986a. Family No. 185: Lethrinidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 595-600.

Smith MM and Heemstra PC. 1986. Smiths’ Sea fishes. Macmillan, Johannesburg.

Smith MM and Smith JLB. 1986. Family No. 183: Sparidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 580-594.

Smith MM and Smith JLB. 1986a. Family No. 222: Mugilidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 714-720.

Smith-Vaniz WF. 1986. Family No. 210: Carangidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 638-661.

Strydom NA and d'Hotman BD. 2005. Estuary-dependence of larval fishes in a non-estuary associated South African surf zone: evidence of continuity of surf assemblages. Estuarine, Coastal and Shelf Science 63: 101-108.

Thompson EF, Strydom NA and Hecht T. 2007. Larval development of Dagetichthys marginatus (Soleidae), obtained from hormone-induced spawning under artificially reared conditions. Scientia Marina 71: 421-428.

Tighe KA and Keene MJ 1984. Zeiformes: development and relationships. Pp393-398. In: ASIH Special Publication 1. (see Moser et al 1984).

Uchida K, Imia S, Mito S, Fujita S, Ueno M, Shojima Y, Semta T, Tahuka M and Dôtu Y. 1958. Studies on the eggs, larvae and juvenile of Japanese fishes. Series 1. Second Laboratory of Fisheries Biology, Fisheries Department, Faculty of Agriculture. Kyushu University pp 1-89 plus 86 plates (in Japanese).

Vachon J, Desoutter M and Chapleau F. 2005. Solea bleekeri Boulenger, 1898, a junior synonym of Pagusa nasuta (Pallas, 1814), with the recognition and redescription of Solea turbynei Gilchrist, 1904 (Pleuronectiformes, Soleidae). Cybium 29: 314-319.

Van der Elst RP. 1981. A guide to the common sea fishes of Southern Africa. Struik, Cape Town. Pp 1-367.

Van der Elst RP. 1988. Shelf Ichthyofauna of Natal. In: Coastal Ocean Studies off Natal, South Africa. E.H. Schumann (ed.) Berlin. Springer Verlag, Chapter 9: pp 209-225.

Weerts SP. 2002. Habitat utilisation by juvenile fishes in Mhlathuze estuary and Richards Bay harbour. MSc thesis, University of Zululand. Pp 1-211.

Whitehead PJP and Wongratana T. 1986. Family No. 55: Engraulidae. In: Smith & Heemstra (Eds.) Smiths’ Sea Fishes: pp. 204-207.

Whitfield AK. 1990. Life-history styles of fishes in South African estuaries. Environmental Biology of Fishes 28: 295-308.

Whitfield AK. 1994. An estuary-association classification for the fishes of southern Africa. South African Journal of Science 90: 411-419.

Wongratana T, Munroe TA and Nizinski MS. 1999. Engraulidae. In: Carpenter KE and Niem VH. (Eds.) FAO species identification guide for fisheries purposes. The living marine resources of the Western Central Pacific. Volume 3. Batoid fishes, chimaeras and bony fishes Part 1(Elopidae to Linophrynidae). Rome FOA. pp. 1698-1753.

Notes on fish eggs, and the species data sheets.

The age of eggs in the samples.

There are 5 major types:-

i)                   eggs >2mm diameter, which can take 8-10 days to hatch, depending on species, lapsed incubation time when collected, and ambient temperatures

ii)                 eggs of 1.5-2mm which might take 48-96 hours to hatch

iii)               eggs ca 1-1.5mm which are usually 20-24 hours into incubation when examined. They were spawned the evening before collection, and will usually hatch by the next evening ie ca 48 hours (or on the evening of collection if spawned two evenings before collection)

iv)               small eggs around 650-800µm, spawned the evening before, and hatching in the bucket if left too long. Incubation time 24-28 hours

v)                 fresh eggs, still in the blastula stage. These are the early morning spawners, if collected during the day. In the Park Rynie area, the species known to be in this group are mostly sparids, including Diplodus sargus (LIIIC1), Chrysoblephus puniceus (LIIID5), Cheimerius nufar (LIIID7)  and Pagellus natalensis, but also Cynoglossus (MIIIA8) and Arnoglossus (LIIIG7), and MIIIA1 and MIIIA2, among others.

Factors affecting egg density.

Apart from the seasonal factors controlling spawning in individual species, and day to day factors such as sudden water temperature changes, the behaviour of the fish themselves also affect the number of eggs collected. Solitary species, such as Synodontidae, are often seen in pairs, but dispersed, and this is reflected in their eggs seldom being found in high numbers in samples. At the other extreme, the highest individual species abundance of eggs were invariably the shoaling species, such as Sardinops, Etrumeus and Scomber, followed by the aggregating species such as Pomadasys striatum, P. olivaceum, and the sparids Diplodus sargus and Pagellus natalensis (see Table 3). These factors should be kept in mind when looking at the data sheets. Note also that by deliberately choosing to sample in the evening at Durban Harbour mouth, any early morning spawners would have been severely under-sampled, since 80-90% of eggs were fresh, and anything older would have been spawned the previous evening ( or early morning), diluted outside the harbour on the outgoing tide, and washed back into the harbour on the next incoming tide.

There is also a collection bias created by the incubation time of the eggs of different species. Thus comparisons of catch-rate should be made with caution unless the eggs have a similar incubation time.

Tips on examining live eggs.

As noted in Methods, I always used reflected light on the dissecting microscope, as I found it preferable for seeing all shades of yellow pigment.

Segmentation within the yolk, when present, is variable. In some species it is evenly distributed through the yolk in small interlinking semicircles, like fine cob-webbing. In others, it can be more symmetrical, in one or two rings near the outer edge of the yolk. In some, for example Trachurus, it is bold and strongly refractory; in others it is more subtle, and requires a light and dark edge under the egg, with reflected light, to see it clearly. I find a small piece of white paper, slid under the watchglass, sitting on a black base, the most convenient.

Yolk surface is another character used, and should not be confused with yolk segmentation. The yolk surface can be smooth or evenly rough, like sandpaper, or the bumps may be a little more separated, variously described here as moon-scaped or goose-pimpled. When examining eggs, be sure they are floating and not touching the bottom of the vial. That way those with oil globules orientate correctly, as seem in the species sheets’ photographs. Some eggs become more dense, and sink to the bottom of the bowl as they develop. Notes on these will be found in the individual sheets.

The oil globule, when present, is always clear (i.e. unsegmented), but it can vary in colour, from colourless, through many shades of amber, to red. The paler shades of amber and pink are best seen on a white background. When there is pigment on the surface of the oil globule, as a rule the yellow pigment is dorsal, and the black pigment ventral, when viewed in the normal resting position, with the oil globule upward.

Size versus age in fish larvae.

For consistency, I have described the larvae by age. But the preference is for size (eg flexion is completed by 6.7mm SL). Thus all larvae and juveniles are measured. So while the discussion will give an event occurring at 15 days, reference to the image will immediately inform the reader of the size of the fish at that point. All measurements are of the live animal unless otherwise noted, and all measurements are notochord length (NL) in preflexion and flexion larvae, and standard length (SL) for postflexion.

Abbreviations and definitions. (see Leis & Rennis for further definitions).

BOLD – Barcode of Life Database

DHM – Durban Harbour Mouth

flexion – a brief stage in larval development, when the posterior tip of the notochord bends upwards, and the caudal (tail) fin takes on the adult appearance

gut length as a percentage of NL – the gut measurement in this context is taken from the same point as the NL, ie the horizontal distance from snout to hind edge of anus.

inshore – a sample collected at sea, about 0.5km offshore, over about 15m water depth.

KZN – KwaZulu-Natal, previously known as Natal, the east coast province of South Africa (see Figure 1)

MPA – Marine Protected Area

neuromasts – small sensory organs, comprising a small raised cone with a sensory filament at the tip. Most newly hatched larvae appear to have about 10 scattered over the head and body. Location appears to have no taxonomic value.

NL  - notochord length (in mm), from snout to notochord tip, in preflexion and flexion larvae (unless otherwise noted, all measurements are of live specimens).

offshore – a surface sample collected at sea, about 5km from shore, over 40-50m of water depth.

ORI – Oceanographic Research Institute, Durban

PC - post-collection, referring to egg or larval age, with collection set at midday, in offshore samples, unless otherwise noted.

PH – post-hatch (unless otherwise noted, all larval age data is PH)

PVS- perivitelline space (see Figure 13)

 

Fig 12a Fig 12b

Figure 13  Egg: 1: pectoral fin buds, 2: chorion, 3: myomeres and myotomes on the developing embryo, 4: tail wrapped around the yolk; 5: narrow perivitelline space; 6: yolk with stellate black pigment spots; 7: unpigmented eyes; 8: otic capsule. Larva. 1: otic capsule: 2: dorsal finfold; 3: myomeres; 4: serrated finfold edge; 5: caudal finfold; 6: notochord tip; 7: ventral finfold; 8: black pigment ventrally along notochord; 9: myotomes; 10: anus; 11: pectoral fin base. Drawings from Brownell 1979.

SL  - standard length, for postflexion juveniles, from snout to the vertical posterior margin of the hypural plate (tail base, excluding caudal rays).

spiny finfold edges-The edge of the dorsal and ventral finfolds sometimes have small spines embedded, giving a serrated appearance. In most cases it starts in the region of a vertical line through the anus, and ends in the taper towards the caudal finfold. In callionymids it is confined further back, in the unpigmented finfold, and extends into and including the caudal finfold. I have seen it on callionymids, mugiloidids, labrids, acanthurids, some pleuronectiformes such as Arnoglossus, and if Mito’s diagnosis is correct, some lophiids. See sheet KIIIB8, for a list of wrasse genera which have spiny finfold edges. It is usually confined to the 2-5 day larva.

tail – posterior to a vertical line through the posterior edge of the anus

trunk – the body between the head and a vertical line through the posterior edge of the anus

Explanation of the coding system used to categorize the eggs.

A.      Eggs in clusters

B.      Eggs oval, oblong, or spindle-shaped. Note this does not include asymmetrical eggs, such as KIIIA6, or eggs with flat spots, such as EIIA1

C.      Eggs with a sculptured, spiky, nippled or reticulated chorion.

D.      Segmented yolk, no oil globule.

E.         "       " , one oil globule, in anterior (bow) position in the yolksac larva.

F.         "       " , one oil globule in posterior (stern) position in yolksac larva.

G.         "       "  , multiple oil globules.

H.      Clear yolk, no oil globule.

K.        "    " , one oil globule in anterior (bow) position in yolksac larva.

L.        "    " , one oil globule in posterior (stern) position in yolksac larva.

M.       "    "  , multiple oil globules.

Size    I       > 1.5mm diameter

                II        1.0 – 1.5mm

                III     < 1.0mm (=1000µm)

D = Etrumeus, Thryssa, some callionymids

E  = carangids, sphyraenids, gerrids, oplegnathids, mullids, haemulids

F = sardine, Coryphaena, stromataeids, sillagids, Acanthistius

G = eels, Gilchristella, Hilsa,

H = fistulariids, some puffers, scorpaenids and lizardfishes

K = anthiinae, some lutjanids,  lethrinids, labrids, cheilodactylids, ambassids, mullids

L = sparids, jawfishes, scorpaenids (most), epinephilids, mugilids, sciaenids, scombrids, acanthurids, pomacanthids, chaetodontids, leiognathids

M = some pleuronectiformes and puffers, Sarda.

By applying this simple key, one can quickly position an egg and early larva, and reduce the search options to a small cluster of species pages, where egg size spans that of the unknown species. Conversely, one can quickly decipher that ABHIIIA1 is an egg originally laid in clusters, oval in shape, with a clear yolk and no oil globule, measuring <1mm. Some species will span two size ranges, so it is necessary to check species sheets either side of the apparent range.

Table 4: The species list (alphabetical, by code):

Code
Order/Family

Lowest ID

Size range in µm
Oil globule
ABH III A1 Scorpaenidae Scorpaenopsis possi & Parascorpaena mossambica
870-1000 x 650-870
---
ABH III A2 Scorpaenidae Unknown
890-940 x 870-920
---
ABK III A1 Carapidae Unknown
1200 x 780
35
BD III A1 Engraulidae Engraulis encrasicolus & Encrasicholina punctifer
1160-1310 x 530-605
---
BF III A1 Engraulidae Stolephorus holodon
1295-1455 x 680-700
120
BH II A1 Unknown Unknown
840-1100 x 790-960
---
BH III A1 Scorpaenidae Unknown
1055 x 910
---
BK III A1 Carapidae Unknown
960-1000 x 790-820
170
BK III A2 Scaridae Scarus rubroviolaceus & S. ghobban
1455 x 500
120
BL I A1 Unknown Unknown
2180 x 2060
120
BL III A1 Scorpaenidae Helicolenus dactylopterus
915-1060 x 770-840
170-190
BL III A2 Scorpaenidae Scorpaenodes ? kelloggi
720-740 x 625-690
120-145
BL III A3 Ogcocephalidae Halieutaea fitzsimonsi 840 190
CD I A1 Regalecidae Regalecus glesne
2740-2950
---
CD I A2 Beloniformes Scomberesox saurus
2280-2920
---
CD III A1 Callionymidae Draculo celetus +
620-760
---
CH I A1 Hemiramphidae Oxyporhamphus micropterus
1870-2110
---
CH I A1A Exocoetidae Unknown
2100-2200
---
CH I A2 Exocoetidae Cheilopogon nigricans
1990-2200
---
CH II A1 Synodontidae Trachinocephalus myops & Synodus indicus
1070-1140
---
CL II A1 Bothidae Unknown
1030-1300
140-150
CL II A2 Myctophidae Inknown
960-1125
190-205
CM I A1 Ostraciidae Tetrosomus concatenatus & Lactoria fornasini
1800-1900
multiple
CM I A2 Soleidae Zebrias cf regani
1510-1680
multiple
CM I A3 Uranoscopidae Uranoscopus archionema
1650-1900
multiple
CM II A1 Macrouridae Caelorinchus or Coryphaenoides sp.
1225
multiple
D I A1 Anguilliformes Gymnothorax spp. +
2330-4020
---
D II A1 Clupeidae Etrumeus teres & E whiteheadi
1160-1320
---
D II A2 Engraulidae Thryssa vitrirostris
985-1060
---
D III A1 Phosichthyidae Vinciguerria nimbaria +
625-720
---
D III A3 Callionymidae Several callionymids but ID uncertain
680-740
---
D III A4 Lophiiformes Unknown but may be callionymid
700-840
---
E I A1 Carangidae Lichia amia
1800-1920
550-600
E I A1A Polyprionidae Polyprion americanus
2090-2110
530
E II A1 Pentacerotidae Pentaceros capensis
1370-1420
310-340
E II A2 Carangidae Seriola dumerili & S. lalandi
1200-1450
240-360
E II A3 Berycidae Centroberyx spinosus
1200-1250
240-265
E II A4 Carangidae Seriola dumerili & S. rivoliana
1000-1075
260-270
E III A1 Sphyraenidae Sphyraena jello
910-980
240-310
E III A1A Kyphosidae Kyphosus bigibbus
910-985
220-250
E III A2 Oplegnathidae Oplegnathus robinsoni
910-940
190
E III A2A Sphyraenidae Sphyraena obtusata
940-1010
240-265
E III A3 Carangidae Unknown
865-890
200-220
E III A3A Sphyraenidae Sphyraena sp.
840-890
215-240
E III A4 Haemulidae Plectorhynchus chubbi
940-960
215
E III A4A Carangidae Carangoides fulvoguttatus
910-940
215
E III A5 Carangidae Trachurus trachurus & Decapterus russelli
850-900
210-220
E III A5B Haemulidae Plectorhinchus flavomaculatus
910
240
E III A6 Carangidae Trachurus trachurus & Decapterus spp.
745-890
205-230
E III A6A Carangidae Elagatis bipinnulata
890
240
E III A7 Carangidae Scomberoides tol
815-840
290-360
E III A7A Unknown Unknown
815
170
E III A7B Trichonotidae Trichonotus sp. n.
790-840
190
E III A8 Berycidae Centroberyx druzhinini
960-1030
190-220
E III A9 Emmelichthyidae Emmelichthys struhsakeri
745-820
170-190
E III B1 Mullidae Upeneus guttatus
720-790
170-180
E III B2 Haemulidae Pomadasys commersonnii
800-840
190-220
E III B3 Haemulidae Pomadasys striatum
750-820
170-190
E III B3A Kuhliidae Kuhlia mugil
850
230
E III B4 Haemulidae Pomadasys olivaceum
750-790
170-205
E III B4A Carangidae Megalaspis cordyla
800
220
E III B5 Priacanthidae Pristigenys niphonia
760
175
E III B6 Carangidae Caranx sem (=C. heberi)
760
210
E III B6A Holocentridae Myripristis berndti
700
190
E III B8 Gerreidae Gerres acinaces
670-770
170-220
F I A1 Anguilliformes ? Ophichthidae
1510-1780
240-290
F I A1 Anguilliformes Unknown
2500-3410
240-190
F I A2 Melanostomiidae Opostomias micripnus
2080-2880
265-290
F I A3 Malacosteidae Malacosteus niger
2400-2450
265-280
F I A4 Echeneidae Remora australis
2210
410
F II A1 Clupeidae Sardinops sagax
1320-1610
145-170
F II A3 Stromataeidae Hyperoglyphe antarctica
1250-1345
360-410
F II A3A Trachichthyidae Gephyroberyx darwinii
1200
240
F II A4 Coryphaenidae Coryphaena hippurus
1320-1560
265-340
F II A5 Stromataeidae Centrolophus niger
1200-1345
360-410
F II A6 Nomeidae Psenes pellucidus
1225
310
F II A7 Oplegnathidae Oplegnathus conwayi
1010-1150
240-265
F II A8 Emmelichthyidae Plagiogeneion rubiginosum
1030-1225
240-275
F II A8A Tetragonuridae Tetragonurus atlanticus
1150
240-265
F II A9 Monodactylidae Monodactylus falciformis
1010-1065
240-270
F II B1 Scorpididae Neoscorpis lithophilus
940-1060
220-290
F III A1 Serranidae Acanthistius joanae
940-985
190-240
F III A2 Dinopercidae Dinoperca petersi
890-960
190-220
F III A2A Dichistiidae Dichistius multifasciatus
865-910
220-250
F III A2B Monodactylidae Monodactylus argenteus
910-1010
240-265
F III A3 Paralichthodidae Paralichthodes algoensis
960-1010
180-220
F III A4 Pomatomidae Pomatomus saltatrix
840-960
220-265
F III A4B Sillaginidae Sillago cf silama
790-865
190-215
F III A5 Chaetodontidae Chaetodon marleyi
790-865
190-215
F III A6 Pomacanthidae Pomacanthus rhomboides
790-890
190-225
F III A7 Sillaginidae Sillago cf sihama
790-865
190-215
F III A9 Sillaginidae Sillago chondropus
745-790
190-215
G I A1 Anguilliformes Unknown
1970-3490
multiple
G I A2 Monocanthidae Monocanthus japonicus
2090-2280
multiple
G II A1 Clupeidae Hilsa kelee
1080-1175
multiple
G III A1 Clupeidae Gilchristella aestuaria
865-890
multiple
H I A1 Exocoetidae Exocoetus monocirrhus
2500-3050
---
H I A2 Trachipteridae Zu cristatus
2065-2185
---
H I A2A Lampridae Lampris cf guttatus
2350
---
H I A3 Fistulariidae Fistularia petimba & F. commersonii
1530-2060
---
H II A1 Pegasidae Eurypegasus draconis
1200-1345
---
H II A2 Scorpaenidae Unknown
1105-1345
---
H II A3 Scorpaenidae Choridactylus natalensis +
1105-1320
---
H II A3A Scorpaenidae Cocotropus monacanthus
1000-1320
---
H II A4 Synodontidae Saurida undosquamis
1020-1130
---
K III A1 Cheilodactylidae Chirodactylus jessicalenorum
910-1030
220-260
K III A2 Labridae Anchichoerops natalensis
840-950
190-220
K III A4 Serranidae Serranus knysnaensis
860-960
145
K III A6 Ophidiidae Brotula multibarbata +
730-1080
155-200
K III A8 Lutjanidae Paracaesio xanthura
820-910
145-170
K III A9 Caesionidae Caesio xanthalytos
840-910
145-190
K III A10 Mullidae Parupeneus fraserorum & P. rubescens
745-890
155-190
K III B1 Lutjanidae Pristipomoides sieboldii & Paracaesio sordida
860-910
110-170
K III B1A Lutjanidae Paracaesio sordida    
K III B2 Cirrhitidae Cyprinocirrhites polyactis
720-840
145-170
K III B2A Cirrhitidae Cirrhitichthys oxycephalus
700-790
95-110
K III B3 Priacanthidae Priacanthus hamrur
720-770
170-180
K III B3A Lethrinidae Gymnocranius cf griseus
625-720
145-190
K III B4 Lethrinidae Lethrinus rubrioperculatus
625-720
145-190
K III B4A Caesionidae Dipterygonotus balteatus
750
145-190
K III B5 Unknown Unknown
720-770
145-185
K III B6 Unknown Unknown
700
120
K III B7 Labridae Unknown
650-700
110-120
K III B8 Labridae Halichoeres lapillus + other wrasse
550-670
120
K III B9 Ambassidae Ambassis dussumieri
620-680
170-190
K III B9A Unknown Unknown
600-670
145-215
K III B9B Ambassidae Ambassis ambassis
625
190
K III B9C Ambassidae Ambassis natalensis
600-670
190
K III C1 Serranidae Pseudanthias gibbosus & P. squamipinnis
630-690
130-150
K III C1A Serranidae Pseudanthias connelli
670
120
K III C2 Serranidae Anthiinae
650
50
K III C3 Labridae Thalassoma trilobatum and other wrasse
550-625
120-145
L I A1 Caristiidae Caristius spp. incl. groenlandicus
1800-1970
530-650
L I A2 Trichiuridae Trichiurus lepturus
1620-1825
385-455
L I A3 Zeidae Zeus faber
1800-2185
310-360
L I A3A Peristediidae Satyrichthys adeni
1620-1990
145-170
L I A4 Bramidae Brama brama
1560-1700
340-410
L I A5 Veliferidae Velifer hypselopterus
1610-1870
145-190
L I A6 Echeneidae Remora brachyptera
1585
340
L II A1 Triacanthodidae Paratriacanthodes retrospinis
1200-1415
205-220
L II A1A Bramidae Taratichthys longipinnis
1345-1440
350-360
L II A2 Pempheridae Pempheris schwenkii
1285-1460
270
L II A3 Aulostomidae Aulostomus chinensis
1270-1440
220-260
L II A4 Triglidae Trigloporus lastoviza africanus
1295-1320
230-325
L II A4A Scombridae Scomberomorus plurilineatus
1250-1320
385-410
L II A4B Luvaridae Luvarus imperialis
1300
265
L II A5 Centriscidae Aeoliscus punctulatus
1150-1320
220-265
L II A5A Tetrarogidae Ablabys binotatus
1300
145-215
L II A6 Sciaenidae Atractoscion aequidens
1125-1250
280-320
L II A6A Ephippidae Tripterodon orbis
1225-1345
240-310
L II A7 Scombridae Scomber japonicus
960-1225
240-300
L II A8 Chiasmodontidae Chiasmodon niger
1150-1345
310-340
L II A9 Macroramphosidae Unknown
1130-1200
240
L II B1 Champsodontidae Champsodon capensis
985-1060
170-200
L II B3 Grammistidae Aulacocephalus temmincki
1090-1200
170-205
L II B4 Sparidae Diplodus hottentotus
960-1060
190-220
L II B5 Triglidae Lepidotrigla faurei
920-1320
230-250
L II B6 Sciaenidae Unbrina robinsoni
1000-1060
220-250
L II B7 Scorpaenidae Apistus carinatus
910-1100
190-220
L II B8 Scorpaenidae Unknown
985-1080
200-220
L II B9 Mugilidae Liza tricuspidens & Myxus capensis
985-1060
340-360
L II B10 Unknown Unknown
1010
240
L II C1 Bramidae Unknown
1080
220
L III A1 Ophidiidae Hoplobrotula gnathopus
960-1010
220
L III A4 Sciaenidae Umbrina canariensis (?+ others )
910-1010
220-240
L III A5 Platycephalidae Cociella heemstrai
910-985
190-240
L III A6 Platycephalidae Platycephalus indicus
940-1030
215-265
L III A7 Mugilidae Liza dumerilii + other mugilids
865-980
290-350
L III A8 Sciaenidae Argyrosomus japonicus
865-985
240-265
L III A9 Unknown Unknown
920
220
L III A10 Dactylopteridae Dactyloptena peterseni
960-1010
260
L III A11 Scombridae Auxis rochei
865-960
220-265
L III A11A Scombridae Euthynnus affinis
865-960
220-240
L III A11B Scombridae Katsuwonus pelamis
950-1050
240-265
L III B1 Sparidae Polysteganus undulosus
960
190
L III B2 Unknown Unknown
950
190
L III B3 Unknown .Unknown
960
170
L III B4 Serranidae Epinephelus malabaricus & E. andersoni
840-950
170-200
L III B7 Mugilidae Myxus capensis + other mugilids
800-920
325-370
L III B9 Sparidae Sarpa salpa
875-985
190-215
L III B10 Sparidae Pachymetopon grande + other sparids
910-985
190-200
L III C1 Sparidae Diplodus capensis + other sparids
890-940
190-200
L III C1B Sparidae Pachymetopon aeneum
890-910
190
L III C1C Sparidae Sparodon durbanensis
890-910
190
L III C2 Unknown Unknown
940-985
170-220
L III C3 Paralichthyidae ?Pseudorhombus sp
840-865
120-145
L III C4 Malacanthidae Malacanthus brevirostris
890-985
170-190
L III C5 Sparidae Pagellus natalensis
790-865
170-190
L III C9 Paralichthyidae Pseudorhombus arsius
840-940
120-170
L III C9C Bothidae Chascanopsetta lugubris
915
145
L III D1 Sparidae Lithognathus mormyrus
910-940
170-190
L III D2 Serranidae Epinephelus poecilonotus & E. rivulatus
815-910
180-190
L III D4 Serranidae Cephalopholis sonnerati
815-865
170-190
L III D5 Sparidae Chrysoblephus puniceus
815-865
180
L III D6 Sparidae Rhabdosargus holubi +
850
180
L III D7 Sparidae Cheimerius nufar
780-950
170-190
L III D8 Zanclidae Zanclus canescens
790-815
190
L III D9 Sparidae Rhabdosargus sarba
790-820
170-190
L III D10 Serranidae Liopropoma cf latifasciatum
840-865
170-180
L III E1 Pinguipdidae Parapercis robinsoni +
750-865
170-190
L III E3 Sciaenidae ? Argyrosomus thorpei
745-790
170-220
L III E3A Sparidae Acanthopagrus vagus
720-770
190-220
L III E4 Caproidae Antigonia rubescens (a cluster of 5 or 6 cryptic spp.)
815-890
180-190
L III E4A Sciaenidae ? Johnius dussumieri
815-890
190-230
L III E5 Macroramphosidae Macroramphosus scolopax
865-985
200-220
L III E7 Acanthuridae Acanthurus mata
670-720
160-180
L III E8 Chaetodontidae Heniochus acuminatus
690-745
170
L III E9 Paralichthyidae Pseudorhombus elevatus
730-810
50-90
L III E10 Acanthuridae Zebrasoma gemmatum
665-770
150-190
L III E11 Sparidae Crenidens crenidens
720-770
180
L III F1 Gadidae Unknown
720-840
145-170
L III F2 Chaetodontidae Chaetodon blackburni, C. dolosus & C. auriga
700-745
170-180
L III F3 Nomeidae Cubiceps pauciradiatus
720-820
190-210
L III F5 Leiognathidae Secutor ruconius
650-720
170-190
L III F6 Acanthuridae (Nasinae) Naso mcdadei, Naso thynnoides & N. brevirostris
670
170
L III F7 Pinguipdidae Parapercis maculata & Parapercis schauinslandii
670-725
145
L III F9 Bothidae Unknown
600-695
120-145
L III G2 Pomacanthidae Centropyge acanthops
615
155
L III G4 Acanthuridae Acanthurus triostegus
625-675
140-160
L III G7 Bothidae Crossorhombus valderostratus
550-610
120
M I A1 Veliferidae Metavelifer multiradiatus
2210-2300
multiple
M I A1A Ostraciidae Ostracion cubicus
1945-2115
multiple
M I A2 Exocoetidae Unknown
1900
multiple
M I A2A Soleidae Unknown
1800
multiple
M I A3 Diodontidae Diodon holocanthus
1610-1870
multiple
M I A4 Pleuronectiformes Unknown
1610
multiple
M II A1 Pleuronectiformes Dagetichthys marginatus
1335-1440
multiple
M II A1 Pleuronectiformes Unknown (Type B)
1405-1415
multiple
M II A1A Molidae Ranzania laevis
1370-1560
multiple
M II A2 Scombridae Sarda orientalis
1200-1370
multiple
M II A2A Soleidae Aseraggodes heemstrai
1150-1320
multiple
M II A3 Platycephalidae Unknown
1100-1225
multiple
M II A4 Bramidae Unknown
1285
multiple
M II A5 Pempheridae Parapriacanthus ransonneti + other pempherids
1100-1200
multiple
M II A6 Cynoglossidae Symphurus sp.
1080-1150
multiple
M III A1 Platycephalidae Rogadius portuguesus
920-970
multiple
M III A2 Platycephalidae Onigocia oligolepis & ?Thysanophrys celebica
890-970
multiple
M III A4 Creediidae Apocreedia vanderhorsti
960-1010
multiple (2)
M III A5 Soleidae Pagusa nasuta (= Solea turbynei,= S. bleekeri)
750-800
multiple
M III A6 Cynoglossidae Cynoglossus lida
770-960
multiple
M III A7 Creediidae Limnichthys nitidus
700-790
multiple
M III A8 Cynoglossidae Unknown
680-700
multiple
M III A9 Unknown Unknown
600
multiple

 

Table 5: The species list (by Order or Family):

Can be seen on the homepage via this link.