Transactions of the Royal Society of South Africa ISSN: 0035-919X (Print) 2154-0098 (Online) Journal homepage: www.tandfonline.com/journals/ttrs20 A preliminary survey of the estuaries on the southeast coast of South Africa, Cape St Francis – Cape Padrone, with particular reference to the fish fauna N.C. James & T.D. Harrison To cite this article: N.C. James & T.D. Harrison (2010) A preliminary survey of the estuaries on the southeast coast of South Africa, Cape St Francis – Cape Padrone, with particular reference to the fish fauna, Transactions of the Royal Society of South Africa, 65:1, 69-84, DOI: 10.1080/00359191003652116 To link to this article: https://doi.org/10.1080/00359191003652116 Published online: 23 Mar 2010. Submit your article to this journal Article views: 1106 View related articles Citing articles: 3 View citing articles Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=ttrs20 Transactions of the Royal Society of South Africa Vol. 65(1), February 2010, 69–84 A preliminary survey of the estuaries on the southeast coast of South Africa, Cape St Francis – Cape Padrone, with particular reference to the fish fauna 1 N.C. James & T.D. Harrison 1 2 South African Environmental Observation Network, Elwandle Node, Private Bag 1015, Grahamstown, 6140 South Africa e-mail: n.james@saiab.ac.za. 2 South African Institute for Aquatic Biodiversity, Private Bag 1015, Grahamstown, 6140 South Africa e-mail: tdharrison@eircom.net A basic ichthyofaunal and physico-chemical survey of the coastal outlets on the southeast coast of South Africa (Cape St Francis – Cape Padrone) was conducted during September and October 1995. Eleven systems have been identified along this stretch of coastline. Four systems (Kromme, Gamtoos, Swartkops and Sundays) are permanently open and provide important habitats for a number of estuarine-resident and marine fish species. Three systems (Seekoei, Kabeljous and Van Stadens) are moderately sized estuaries characterised by intermittently open mouths. The Seekoei Estuary has undergone extreme human disturbance, while the Kabeljous and Van Stadens are in a less disturbed state. All three systems provide nursery habitats for both estuarine-resident and estuarine-associated marine species. The Maitland is a small, shallow sandy system that is typically closed to the sea for extended periods; despite this it was still found to support estuarine-associated species particularly mugilids. The remaining three systems (Bakens, Papkuils and Coega) are all highly altered systems that provide little littoral habitat for either resident or migrant marine species. Keywords: ichthyofauna, estuarine survey, fish habitat, southeast coast, South Africa. INTRODUCTION The South African coastline extends approximately 3000 km from the Orange (Gariep) River (28°38’S, 16°28’E) on the west (Atlantic Ocean) coast to Kosi Bay (26°54’S, 32°53’E) on the east (Indian Ocean) coast. Some 300 coastal outlets have been identified along the coast of South Africa and these range from relatively large, permanently open estuaries to inlets of coastal streams and even dry river beds that only occasionally contain water. Estuaries are typically formed where freshwater from rivers and saltwater from the sea meet and, as such, are unique environments characterised by variations in environmental conditions. Estuaries are amongst the most variable aquatic environments on earth and are known to be more productive than adjacent freshwater and marine environments (Woodwell et al., 1973; Haedrich & Hall, 1976). Estuaries are also sheltered environments and often serve as important nursery areas for various fish species (Dando, 1984; Wallace et al., 1984). After reviewing the available scientific information on South African estuarine systems Whitfield (2000) found that, of the 258 systems included in the review, the state of information on 68% was either ‘nil’ or ‘poor ’; the state of information on 22% was classified as ‘moderate’, while only 10% were regarded as ‘good’ or ‘excellent’. This paper reports on ichthyofaunal surveys of estuaries between Cape St Francis and Cape Padrone on the southeast coast of South Africa conducted during 1995 and describes physico-chemical variables, presents basic fish community data, and provides an appraisal of the nursery potential ISSN 0035-919X print © 2010 Royal Society of South Africa DOI: 10.1080/00359191003652116 http://www.informaworld.com Published online 23 Mar 2010 of these systems for fishes. This paper is a continuation of the published series on the fish assemblages of South African estuaries (Harrison 1997a,b; 1998; 1999a,b; James & Harrison, 2008, 2009). STUDY AREA The coastline in the vicinity of Port Elizabeth is dominated by westerly and southwesterly winds throughout the year (Stone et al., 1998). This coastline borders the southwestern Indian Ocean and is influenced by the warm, south-flowing Agulhas Current (Heydorn, et al., 1978). Eleven river outlets intersect the coast in this geographical sector (Figure 1). Rainfall along the coastline west of Port Elizabeth occurs throughout the year with maximum falls usually experienced during spring and autumn (Kopke, 1988). The coastline that extends from Port Elizabeth to Port Alfred tends to receive most of its rain in winter (Kopke, 1988). MATERIALS AND METHODS The estuaries between Cape St Francis and Cape Padrone were sampled during September and October 1995. Each system was sampled once and took 1–3 days to survey, depending on the size of the system. Physico-chemical During each survey, selected physico-chemical parameters were measured at various sites within each system. Water depth and transparency were measured using a 20 cm diameter 70 Transactions of the Royal Society of South Africa Vol. 65(1): 69–84, 2010 Figure 1. The locality of study estuaries between Cape St Francis and Cape Padrone on the southeast coast of South Africa. Secchi disc attached to a weighted shot line graduated at 10 cm intervals. Temperature (°C), salinity (‰), conductivity (mS cm–1), pH and dissolved oxygen (mg l–1) were measured using a Horiba U-10 Water Quality Checker. Where water depth permitted (usually >0.5 m), both surface and bottom waters were measured. The mouth state of each system at the time of sampling was also noted. Ichthyofauna The ichthyofauna of each estuary was sampled using a 30 m × 1.7 m × 15 mm bar mesh seine net with a 5 mm bar mesh purse and a fleet of gill nets. The gill nets were either 10 m or 20 m in length and 1.7 m in depth and consisted of three equal sections of 45 mm, 75 mm and 100 mm stretch meshes. Seine netting was carried out during daylight hours in shallow (<1.5 m deep), unobstructed areas with gently sloping banks. Fish caught were identified and measured to the nearest millimetre standard length (SL) before being released. Where large catches of a species were made a sub-sample was kept and returned to the laboratory where they were identified, measured (mm SL) and weighed to the nearest 1.0 g. The remaining specimens were counted and the batch weighed. Specimens that could not be identified were also kept and identified in the laboratory by reference to Smith & Heemstra (1991) and Skelton (1993). Taxonomic identities of certain species were adjusted using information provided in Heemstra & Heemstra (2004). The total species composition, by number and mass, was calculated for each system. The relative biomass contribution of each species was calculated using actual recorded masses as well as masses derived from length–mass relationships presented in Harrison (2001). Where appropriate, the length frequency distribution (in 10 mm size classes) was calculated for the most abundant species within each system. Using information from Whitfield (1998) the species recorded were divided into four estuarine association categories: freshwater, estuarine-resident, estuary-associated marine and marine species. The relative contribution made by each category to the total ichthyofaunal assemblage of each system was calculated in terms of number of species, relative abundance and relative mass. RESULTS AND DISCUSSION Kromme The Kromme Estuary (34°09’S, 24°51’E) is situated approximately 55 km west of Port Elizabeth and is one of the largest estuaries in the area (Strydom & Whitfield, 2000). The Kromme River is about 95 km long and drains a catchment area of James & Harrison: Survey of the estuaries on southeast coast of South Africa, Cape St Francis – Cape Padrone 71 Table 1. Physico-chemical parameters measured in southeast coast estuaries (Cape St Francis – Cape Padrone) during September/October 1995 (S, surface; B, bottom; 999, Secchi disc visible on the bed). System Site Depth (m) Temperature (°C) Salinity (‰) Conductivity (mS cm–1) pH Dissolved oxygen –1 (mg l ) S B S B S B S B S B Secchi depth (m) Kromme 1 2 3 4 5 6 7 8 9 0.9 3.0 3.0 1.4 2.9 1.2 3.2 3.2 3.4 17.9 17.8 18.3 18.2 18.7 19.6 19.9 19.8 20.6 17.9 17.9 18.3 18.2 18.4 19.1 18.9 19.6 20.1 32.5 32.2 32.0 31.4 30.6 28.4 28.6 26.9 27.7 32.8 32.7 32.0 31.5 30.7 28.4 29.2 28.0 28.5 49.9 49.7 49.0 48.3 46.9 43.8 44.1 42.0 42.9 50.1 50.1 49.0 48.4 47.4 44.5 45.3 43.7 44.4 7.8 7.8 7.8 7.8 7.8 7.5 7.8 7.8 7.8 7.9 7.9 7.8 7.8 7.8 7.5 7.8 7.8 7.7 7.7 7.5 6.9 6.4 6.6 5.2 6.5 6.6 6.6 7.7 7.7 7.0 6.5 6.5 4.3 6.3 6.5 5.0 999 1.3 0.9 0.9 0.9 0.6 1.7 2.1 1.9 Seekoei 1 2 3 4 1.0 0.5 1.6 1.2 17.6 20.4 18.7 20.8 17.7 20.4 18.7 19.8 9.0 6.9 4.6 3.8 9.3 6.9 4.6 3.9 15.4 12.0 8.4 7.1 15.9 12.1 8.5 7.3 8.3 8.8 8.4 9.2 8.2 8.8 8.5 9.4 8.0 12.6 8.9 11.5 9.7 12.4 8.9 12.9 999 999 0.6 0.9 Kabeljous 1 2 3 1.0 0.9 1.1 17.2 17.5 17.9 17.3 17.6 18.4 17.1 16.5 15.2 16.9 16.5 16.5 27.5 26.7 24.7 27.4 26.9 26.8 8.0 7.9 8.1 8.0 7.9 8.0 6.2 4.6 7.1 5.0 4.1 5.7 999 999 999 Gamtoos 1 2 3 4 5 6 7 8 1.3 1.1 2.6 2.6 1.9 1.1 1.1 1.0 18.5 18.6 18.6 20.0 20.2 19.4 19.7 19.6 18.5 18.6 18.0 19.2 19.4 19.4 19.4 19.4 31.4 29.6 18.8 22.8 15.0 10.7 7.4 3.1 31.4 29.8 23.8 28.6 28.0 24.5 8.8 3.1 48.2 45.7 30.3 35.9 22.2 16.3 12.9 5.86 48.2 45.9 37.5 44.5 43.8 39.0 15.2 5.92 7.8 8.0 8.1 8.1 8.3 8.4 8.3 8.2 7.9 8.0 8.0 7.9 7.9 7.6 8.0 8.2 7.8 7.6 8.4 8.4 9.5 10.3 11.6 9.8 7.8 7.7 8.1 7.2 6.3 3.3 5.8 9.7 0.3 1.0 0.9 1.0 0.7 0.6 0.5 0.6 Van Stadens 1 2 3 4 2.6 3.3 1.4 1.5 19.5 20.5 21.4 22.0 18.5 20.6 21.4 21.6 15.0 15.0 15.2 10.5 15.1 15.3 15.1 15.2 24.5 24.6 24.7 16.2 24.8 25.1 24.9 25.1 7.8 7.8 7.6 7.3 7.8 7.7 7.6 7.3 7.5 6.9 6.0 5.3 7.2 7.0 5.7 3.1 999 2.5 999 999 Maitland 1 0.3 22.7 Bakens 1 2 3 1.0 0.2 0.2 19.2 19.0 20.5 19.2 29.8 9.6 0.7 31.7 46.2 20.1 1.6 48.6 7.8 7.5 8.1 6.8 999 999 999 18.2 18.4 18.5 18.5 18.7 19.1 32.7 32.9 31.5 30.2 27.0 25.2 32.7 32.9 31.6 30.2 28.6 27.0 50.0 50.3 48.3 46.5 41.9 39.5 50.0 50.3 48.4 46.5 44.3 42.0 7.9 7.9 7.7 7.7 7.5 7.4 Papkuils 1 0.1 24.0 Swartkops 1 2 3 4 5 6 2.3 0.6 2.4 2.5 2.0 3.5 18.2 18.4 18.5 18.5 18.8 19.2 1.3 2.6 9.7 8.4 15.7 12.9 7.9 6.2 9.1 15.9 7.9 7.9 7.8 7.7 7.6 7.5 6.8 7.7 6.3 6.1 5.6 5.0 7.4 999 3.4 999 6.9 7.8 6.5 6.2 5.3 5.0 1.1 999 1.0 1.2 1.3 1.6 Coega 1 0.7 13.4 13.3 47.2 47.2 61.6 61.7 8.2 8.2 7.2 7.1 999 Sundays 1 2 3 4 5 6 7 1.2 3.5 2.1 3.5 1.9 2.5 2.2 19.1 20.0 21.2 21.9 21.5 22.1 21.9 18.9 19.1 19.6 19.8 20.2 20.6 20.4 32.8 22.6 15.4 13.9 10.4 6.5 3.4 32.8 24.3 26.6 24.8 21.7 15.1 3.2 50.2 35.6 25.0 22.7 17.6 11.5 6.4 50.1 38.0 42.1 39.0 34.7 24.8 6.0 8.1 8.0 8.1 8.1 8.2 8.4 8.5 8.1 8.0 7.8 7.9 7.9 8.1 8.5 8.7 7.9 8.3 9.0 8.8 8.9 9.9 8.9 6.6 4.9 5.5 5.4 5.9 8.1 0.5 0.7 0.6 0.5 0.4 0.4 0.4 approximately 936 km2 (Reddering & Esterhuysen, 1983). Numerous tributaries enter the Kromme, with the main tributary being the Geelhoutboom that flows into the estuary about 9 km upstream of the mouth (Bickerton & Pierce, 1988). There are two large dams in the catchment and these severely restrict freshwater input into the estuary. The Kromrivier Dam, which was completed in 1943, is situated approximately 35 km from the head of the estuary, while the Mpofu Dam, completed in 1982, is situated 4 km from the tidal head. Together these dams have the capacity to store about 133% of the mean annual runoff of the Kromme River catchment (Scharler & Baird, 2000). In addition, there are also numerous small farm dams 72 Transactions of the Royal Society of South Africa within the catchment that further restrict the flow of freshwater into the estuary (Bickerton & Pierce, 1988). The upper part of the estuary is narrow and is bounded on either side by rocky cliffs but nearer the sea the banks become less steep and the system widens before discharging into St Francis Bay (Marais, 1983a). A well-developed flood-tidal delta is present in the lower reaches, and these sand banks are exposed at low tide (Bickerton & Pierce, 1988). The Kromme Estuary is a popular recreational area; a marina development is situated near the mouth and numerous holiday houses have been built further upstream. Two road bridges cross the estuary approximately 3 km and 15 km from the mouth, respectively. According to Whitfield (2000) the state of information on the Kromme is regarded as excellent. Physico-chemical Physico-chemical parameters were measured at eight sites from the mouth to the head of the estuary; one site (Site 6) was located in the inlet of the Geelhoutboom tributary. The mouth of the Kromme Estuary is permanently open; the presence of a flood-tidal delta in the lower reaches of the system, together with the low river inflow, indicates that tidal currents serve to maintain a connection with the sea. The channel depth during this survey ranged from 0.9 m near the mouth to 3.4 m recorded at the uppermost site (Table 1). Bickerton & Pierce (1988) reported similar water depths of between 1.5 and 3.5 m in the system. Surface water temperatures during this survey ranged between 17.8°C recorded in the lower reaches and 20.6°C at the uppermost site; bottom water temperatures showed a similar pattern and ranged between 17.9 and 20.1°C (Table 1). Overall, surface values were slightly higher than those at the bottom and this is probably due to solar radiation warming the surface waters. The influence of cooler marine waters also probably accounted for the horizontal thermal gradient recorded. Day (1981) reports a seasonal temperature range of between 14.0 and 24.0°C for the Kromme Estuary. According to Bickerton & Pierce (1988) surface water temperatures in the system range from 12.0°C in winter to 28.0°C in summer. Average summer temperatures recorded by Scharler et al. (1997) measured between 20.0 and 24.0°C, while winter temperatures averaged 17.0–18.0°C. Both surface and bottom salinities recorded during this survey exhibited a horizontal salinity gradient and decreased from the mouth upstream (Table 1). Surface salinities ranged between 32.5 and 26.9‰; bottom salinities were slightly higher and measured between 32.8 and 28.0‰. According to Hanekon & Baird (1984) salinities in the Kromme Estuary are normally about 33.0‰ but can decline to as low as 1.0‰ during winter floods. Extensive water abstraction in the catchment, however, has resulted in high salinities in the estuary with occasional incidences of hypersaline conditions developing in the upper reaches (Bickerton & Pierce, 1988; Scharler & Baird, 2000). Average salinities in the Kromme vary between 26.0 and 33.0‰ (Emmerson & Erasmus, 1987; Scharler et al., 1997). Salinties above 28.0‰ have been recorded throughout the estuary, even following heavy rain and the release of freshwater from the Mpofu Dam (Bickerton & Pierce, 1988; Heymans & Baird, 1995; Scharler & Baird, 2000; Strydom & Whitfield, 2000). The pH of the water in the Kromme Estuary ranged from 7.5 to 7.8 (Table 1). Heymans & Baird (1995) recorded an annual mean pH of 8.0 reflecting the strong marine influence. Hecht (1973) found that the pH of Kromme Estuary water ranged between 7.9 and 8.4. The dissolved oxygen of both the surface and bottom waters decreased from the mouth upstream (Table 1). Surface water dissolved oxygen concentrations Vol. 65(1): 69–84, 2010 ranged from 6.4 to 7.7 mg l–1 while those at the bottom ranged from 5.0 to 7.7 mg l–1. Surface water dissolved oxygen concentrations, particularly those above the Geelhoutboom tributary were also slightly higher than those at the bottom and this is probably a reflection of salinity stratification and lack of mixing. Hecht (1973) found little variation between surface and bottom waters, with surface values ranging from 5.7 to 7.9 mg l–1. Overall, the waters of the Kromme Estuary are well oxygenated with dissolved oxygen values generally exceeding 6.0 mg l–1 (Emmerson & Erasmus, 1987; Bickerton & Pierce, 1988; Heymans & Baird, 1995; Scharler et al., 1997). The waters in the estuary were relatively clear with Secchi disc values generally exceeding 0.9 m (Table 1). This is probably a result of the strong marine influence in the system. Other workers also found the waters in Kromme Estuary to be fairly clear with mean Secchi disc values measuring between 0.8 and 1.8 m (Marais, 1984; Bickerton & Pierce, 1988; Scharler et al., 1997). Ichthyofauna Eighteen seine net hauls and nine gill nets captured a total of 4101 individuals representing 30 species from 15 families. Mugilidae (6 species), Gobiidae (5 species) and Sparidae (4 species) dominated the taxa. Atherina breviceps was the most abundant species caught comprising 41.3% of the total catch followed by Rhabdosargus holubi (14.7%), Gilchristella aestuaria (13.9%), Liza dumerili (6.2%), Glossogobius callidus (4.8%), Liza richardsonii (4.2%), juvenile mugilids (3.1%), Psammogobius knysnaensis (2.7%), Lithognathus lithognathus (2.1%) and Liza tricuspidens (1.1%). Together these species comprised 94% of the total catch numerically (Table 2). A total species mass of over 95 kg was caught. Liza richardsonii was the dominant species comprising 28.7% of the overall biomass, followed by Lichia amia (23.1%), Pomadasys commersonnii (11.2%), Liza tricuspidens (9.8%), Galeichthys feliceps (6.1%), Mugil cephalus (4.8%), L. dumerili (3.9%), R. holubi (3.8%), A. breviceps (1.9%), Argyrosomus japonicus (1.8%) and Platycephalus indicus (1.1%) (Table 3). A total of 36 species has been recorded in the Kromme Estuary from various surveys using either gill nets and/or seine netting (Baird et al. 1981; Marais, 1983a; Bickerton & Pierce, 1988; Hanekom & Baird, 1988). Twenty-four species (67%) were common to those recorded during this survey. Many of the important species reported were also similar to those that dominated during this survey. Dominant taxa either numerically and/or in terms of biomass included A. japonicus, A. breviceps, Caffrogobius gilchristi, G. feliceps, G. aestuaria, G. callidus, L. amia, L. dumerili, L. richardsonii, L. tricuspidens, Monodactylus falciformis, P. commersonnii, P. knysnaensis and R. holubi (Marais, 1983a; Hanekom & Baird, 1984; Bickerton & Pierce, 1988). Of the species collected in the Kromme during this survey, nine were species that are able to live and breed in estuaries; these included A. breviceps, Caffrogobius nudiceps, C. gilchristi, Caffrogobius natalensis, Clinus superciliosus, G. aestuaria, G. callidus, P. knysnaensis and Syngnathus temminckii. Atherina breviceps captured during this survey measured between 29 and 58 mm; the majority of these were mature individuals, which is estimated to occur at a size of 40 mm SL (Ratte, 1989). Psammogobius knysnaensis ranged in size from 17 to 48 mm and most of these were also mature individuals in the 30–40 mm size class; P. knysnaensis matures at a length of approximately 30 mm SL (Bennett, 1989). The remaining species captured during this survey were estuarine-associated marine species whose juveniles are dependent on estuaries to varying degrees. These included A. japonicus, Diplodus hottentotus, Diplodus capensis, G. feliceps, Heteromycteris capensis, L. amia, L. lithognathus, L. dumerili, Liza Table 2. Relative abundance (%n) of fishes captured in all net types (seine (S) and gill (G) nets) in southeast coast estuaries (Cape St Francis – Sundays) during September/October 1995. Species Kromme S G Seekoei %n S G Kabeljous %n S G %n Gamtoos S G Van Stadens %n S G %n Maitland S G Bakens %n S G Swartkops %n S G %n 8 0 0.2 0 3 0.1 51 0 1.1 S G Sundays %n 0.7 57 0 1.6 2.3 95 0 2.7 0.4 1 0 0.0 0 1 0.0 0.0 Atherina breviceps 1695 0 41.3 Caffrogobius gilchristi 12 0 0.3 110 0 Caffrogobius natalensis 12 0 0.3 19 0 Caffrogobius nudiceps 13 0 0.3 6 0 0.1 Clinus superciliosus 25 0 0.6 4 0 0.1 Diplodus sargus capensis 10 0 0.2 29 0 0.6 Diplodus cervinus hottentotus 1 0 0.0 21.8 1529 0 68.0 0.6 219 0 5.1 31 0 0.7 141 0 8.2 4 0 4.8 1895 0 96.8 Carcharias taurus 23 Elops machnata Galeichthys feliceps 0 12 0.3 Gilchristella aestuaria 570 0 13.9 345 0 28.9 Glossogobius callidus 198 0 4.8 31 0 2.6 1 0 0.0 0 0.5 1 0 0.1 1 0 1.2 0.1 0 23 0 109 2.6 0 16 0.3 0 8 0.2 1136 0 26.6 1 0 0.1 1389 0 28.7 30 0 1.5 1160 0 32.7 8 0 0.2 17 0 1.0 69 0 1.4 1 0 0.1 6 0 0.2 2 0 0.0 29 0 0.6 5 0 0.3 20 0 0.6 0 5 0.1 0 1.0 2 0 0.2 29 0 0.7 25 0.6 0 3 0.3 0 3 0.1 0 3 0.1 0 4 0.2 Lithognathus lithognathus 85 1 2.1 190 7 16.5 118 2 5.3 106 1 2.5 193 4 11.5 Liza dumerilii 245 9 6.2 10 10 1.7 676 14 16.2 15 0 0.9 Liza macrolepis 1 0 0.0 Liza richardsonii 99 75 4.2 1 10 0.9 Liza tricuspidens 26 18 1.1 43 Mugil cephalus 17 7 0.6 5 32 Juvenile mugilids 128 0 3.1 259 0 Myxus capensis 0 3 0.1 Monodactylus falciformis 9 4 0.3 184 70 11.3 483 36 12.1 1 47 2.8 4 0 0.2 1 14 0.4 0 3 0.2 3.1 21 4 1.1 137 3 3.3 38 0 2.2 21.7 261 0 11.6 340 0 8.0 922 0 1 5 0.3 25 6 0.7 79 0 1 4 0.2 1 2 0.1 5 3 0.5 5 0.4 0.6 5 0 0 0.0 0 40 Oreochromis mossambicus 0.1 0.1 Lichia amia 0.2 0 0 4 Heteromycteris capensis 2 4 1 0 Hemiramphus far 0 0.0 0 14 2 0 25 %n 1 0 260 0 G 12 Amblyrhynchotes honckenii Argyrosomus japonicus S 40 2 0 0 6.8 0.3 0 51.2 8 0 9.5 83 0 1.7 92 1 2.6 3 0 3.6 232 3 4.9 194 4 5.6 3 0 3.6 284 7 6.0 211 18 6.5 3 13 0.3 2 35 1.0 139 19 4.5 618 0 17.4 0 1 0.0 15 1 0.5 516 0 88.2 4 0 4.8 10 2 0.2 53.7 2 0 0.3 4 0 4.8 663 0 13.7 4.6 16 0 2.7 4 0 12 5 0 0 0.6 0.3 0.1 Platycephalus indicus 0 1 0.0 0 1 0.0 3 0 0.1 1 0 0.0 Pomadasys commersonnii 17 15 0.8 38 4 1.0 750 6 15.6 131 11 4.0 Pomatomus saltatrix 0 4 0.1 Psammogobius knysnaensis 112 0 2.7 Rhabdosargus globiceps 5 0 0.1 Rhabdosargus holubi 599 2 14.7 Sarpa salpa 2 0 0.0 Solea turbynei 1 0 0.0 Syngnathus temminckii 1 0 0.0 Total individuals 3923 178 1119 73 2159 89 4047 225 1657 61 585 0 84 0 4781 60 1957 Total nets 18 9 11 5 13 5 16 15 11 5 5 0 3 0 19 14 5 9 7 0 4 0.8 0.9 3 36 0 1 0.1 1.6 1 1 0.0 52 0 1.2 76 0 1.8 650 2 15.3 1 0 0.1 242 0 14.1 1 0 0.1 1 8 0 0 0.2 1.4 11 3 0 0 Sillago sihama 15 0 0.4 13.1 3.6 2 0 0.0 53 0 1.1 127 0 2.6 824 0 17.0 18 0 0.4 1 0 0.0 8 0 0.2 5 4 0 0 0.3 0.2 31 0 0.9 50 0 1.4 505 1 14.3 66 0 1.9 1 0 0.0 0 3413 137 0 16 14 James & Harrison: Survey of the estuaries on southeast coast of South Africa, Cape St Francis – Cape Padrone Ambassis dussumieri Coega 73 74 Table 3. Relative biomass ( %g) of fishes captured in all net types (seine (S) and gill (G) nets) in southeast coast estuaries (Cape St Francis – Sundays) during September/October 1995. Species Kromme S G Seekoei %g S G Kabeljous %g S G Gamtoos %g S G Van Stadens %g S G Maitland %g S G Bakens %g S G Swartkops %g Ambassis dussumieri Coega S G %g 2.1 0 0.0 0 2215 4.3 S G Sundays %g Amblyrhynchotes honckenii 0 1766 1.8 Atherina breviceps 1848 0 1.9 Caffrogobius gilchristi 17.5 0 0.0 Argyrosomus japonicus 0 261.5 0 1.0 1395 0 3.0 30545 17.9 189.3 0 0.1 28.9 0 0.0 298.7 0 0.9 5.2 0 2.4 S G %g 9.9 0 0.0 428.5 9364.4 7.3 79.3 0 0.2 130.0 0 0.1 125.5 0 0.2 1597 0 91.2 126.2 0 0.1 0.3 0 0.0 Caffrogobius natalensis 9 0 0.0 16.2 0 0.0 Caffrogobius nudiceps 23 0 0.0 16.0 0 0.0 Clinus superciliosus 39.4 0 0.0 8.8 0 0.0 Diplodus sargus capensis 1.6 0 0.0 11.7 0 0.0 Diplodus cervinus hottentotus 0.4 0 0.0 Galeichthys feliceps 0 5884 6.1 Gilchristella aestuaria 256.6 0 0.3 144.7 0 0.5 747.5 0 0.4 1.2 0 0.0 1285.6 0 2.5 20 0 1.1 1054 0 0.8 Glossogobius callidus 208.8 0 0.2 45.5 0 0.2 19.9 0 0.0 25.4 0 0.1 79.2 0 0.2 5.1 0 0.3 8.26 0 0.0 0.0 Heteromycteris capensis 19.4 0 0.0 1.8 0 0.0 17.3 0 0.0 2.2 0 0.1 11.8 0 0.0 Lichia amia 0 22206 23.1 0 1375 5.1 0 4999 10.8 0 863 0.5 0 4047 12.2 85.0 1390 3.2 1857 1237 1.8 55.6 219 0.8 7520 2056 5.6 52.0 0 0.2 0 Carcharias taurus 6.1 13733 0 0.9 0 0.0 0.0 0.2 0 0.0 0.4 0 8.1 0.2 0 62760 36.9 Hemiramphus far Lithognathus lithognathus 207.0 49 0.3 56.1 3614 13.5 Liza dumerilii 2604 1115 3.9 1187 1291 9.1 Liza macrolepis 612.9 0.6 3.4 10.2 0 2.6 0 1.2 115.6 0 0.2 157.9 50 0.2 0 13.1 3300 417 7.2 2718 641 2.5 119 0 56.0 2717 2701 10.5 20.7 3849 7.5 4.8 12.5 13886 51.1 16.3 0.3 28.4 453.9 Myxus capensis 0 952 1.0 Monodactylus falciformis 6.1 198 0.2 0 1076 1.1 0 979 0.6 26.0 0 0.1 9.6 526.8 10215 11.2 82.4 2459 1.5 721.8 4754 10.6 360.3 Pomadasys commersonnii 0 438 0.5 Psammogobius knysnaensis 92.2 0 0.1 Rhabdosargus globiceps 18.8 0 0.0 Rhabdosargus holubi 3491.1 166 3.8 1.0 0 0.0 Solea turbynei 2.3 0 0.0 Syngnathus temminckii 1.5 0 0.0 Total mass 13452 82484 Sarpa salpa 7.3 13.7 165.8 0 170 0.1 1.2 1643 2359 2.4 13.6 0 0.0 357.3 0 95.3 18.8 0 8.8 44.7 961 1.9 1.0 590.1 0 0.3 1492.4 0 4.5 0.2 0 0.0 0.8 0 0.4 1516 0 2.9 0.2 2344 5.0 5.5 2476 1.5 91.3 0 0.3 3.8 0 1.0 0.1 233 0.8 144 0.1 0.9 79 0.2 10.3 6.7 0 0 43 0.5 0.0 0.1 1.69 98 0.1 62.1 0 0.0 125.2 0 0.1 6067 74 3.6 1.2 0 0.0 116.4 0 0.4 0.5 0 0.0 11.4 1.3 0.9 0 0 0.4 0.2 9.2 16.4 0 0 Sillago sihama Total nets 18 9 24.73 1879 25149 6466 39964 11 13 5 5 0 25169 145024 16 15 0.0 2528 30737 375 0 212 0 11 5 0 3 0 5 4.3 7.7 0 5.8 0 0.0 0 0.1 267.8 0 0.5 2614 0 5.0 14.1 0 0.0 2.3 0 0.0 19.7 0 0.0 19 14 3.8 0 0 6.1 0.2 0.0 52.5 13098 38725 106 15.4 2.4 0 0 1751 0 5 0 0.9 0.1 1991 6088 6.0 14.9 33304 24.8 145.7 9300 7.0 1005 0 0 329 0.7 0.2 34.0 60 0.1 0 0.0 5085 4.0 34.4 0 0.0 195.8 0 0.1 2595 64.7 2.0 134.1 0 0.1 0.9 0 0.0 11170 123393 16 14 Vol. 65(1): 69–84, 2010 Pomatomus saltatrix 1994 5204 11.2 0.8 34.9 4090 3.0 27.8 0 0.0 0 4.6 645.3 3999 0 3.0 2369 303.9 0 10.3 0.0 0 Juvenile mugilids Platycephalus indicus 3425 0 Mugil cephalus 0.2 0 0 0 9.8 59 7.0 6.7 17.6 47017 2013 25533 28.7 0 99.0 11880 2.3 0 18.1 502.8 8887 0.1 6081 13361 11.4 378.4 22967 70.2 16.2 0 5.8 9370 0.0 0.0 Liza tricuspidens 0 4481 25751 65.1 8.5 0 23.3 0 0 0 Liza richardsonii Oreochromis mossambicus 2760 12.3 12089 4.3 0.4 Transactions of the Royal Society of South Africa 0 Elops machnata 0 8000 5.9 James & Harrison: Survey of the estuaries on southeast coast of South Africa, Cape St Francis – Cape Padrone macrolepis, L. richardsonii, L. tricuspidens, M. falciformis, M. cephalus, Myxus capensis, P. indicus, P. commersonnii, Pomatomus saltatrix, Rhabdosargus globiceps, R. holubi, Sarpa salpa and Solea turbynei. Lithognathus lithognathus ranged in size from 18 to 130 mm SL. Most of the individuals were newly recruited and fell in the 20–30 mm size class; a few larger individuals (>80 mm) were also present. Lithognathus lithognathus enters estuaries along the southeastern, southern and southwestern Cape coasts at sizes below 50 mm TL (Bennett, 1993). The predominance of small juveniles indicates recent recruitment into the system. Liza dumerili ranged in size from 45 to 234 mm with two size cohorts (60–70 mm and 170–180 mm) predominating; this also suggests regular recruitment and utilisation of the system. Recruitment of L. dumerili into estuaries on the southern Cape coast occurs at <30 mm TL (Whitfield & Kok, 1992). Liza richardsonii ranged in size between 44 and 340 mm and showed a polymodal length frequency distribution; the first size class comprised individuals of 60–70 mm, the second size class was 120–130 mm and the third size class was 280–290 mm. The range of size cohorts captured indicates regular recruitment and utilisation of the system. Juvenile L. richardsonii recruit into estuaries on the southwestern and southern Cape coasts at about 20 to 50 mm TL (Bennett, 1989; Whitfield & Kok, 1992) and a length of about 75 mm FL is attained after a year (Ratte, 1989). Rhabdosargus holubi individuals ranged in size from 12 to 160 mm with most individuals less than one year old in the 50–60 mm size classes; R. holubi attains a length of approximately 100 mm SL by the end of their first year (Beckley, 1984). The predominance of small individuals indicates that the Kromme serves as an important nursery area for this species. During this survey estuarine-resident species comprised 66.4% of the total catch numerically and 2.6% of the biomass, while estuarine-associated marine species comprised 33.6% of the catch numerically and 97.4% of the biomass. The high proportion of estuarine and marine species in the Kromme Estuary suggests that it serves an important function as an estuarine nursery area for both resident and marine species. This is also supported by the high numbers of mature estuarine-resident species as well as the occurrence of a range of size classes of estuarine-associated marine species. Seekoei The Seekoei (34°05’S, 24°54’E) is situated near the coastal town of Jeffrey’s Bay. The river has a total length of approximately 35 km and a catchment area of 250–502 km2 (Bickerton & Pierce, 1988). The main tributary, the Swartrivier, joins the Seekoei approximately 1 km upstream of the mouth so that the estuary consists of two arms; with the main arm formed by the Seekoei River and the secondary arm by the Swartrivier (Bickerton & Pierce, 1988). Numerous farm dams have been built in the catchment, resulting in reduced river flow to the estuary. The lower reaches of the estuary have been severely modified. The mouth dynamics of the system were changed with the construction of a protective embayment and parking area at the coast, followed by a swimming pool complex (Bickerton & Pierce, 1988). This caused a southerly deflection of the position of the mouth and as a consequence the estuary drained completely when open (Dundas, 1994). In an attempt to retain water in the estuary a causeway, which also carried a road, was built about 700 m upstream of the mouth. This causeway only had a narrow opening in the centre and was extremely restrictive to river flow (Dundas, 1994). Since this survey was undertaken, however, the opening in the causeway has been increased (DWAF, 2006). In addition to the causeway, 75 a concrete canal was constructed in an attempt to fix the position of the mouth at its historical location. This canal was artificially maintained and permitted limited tidal exchange to take place; this practice has stopped, however, and the canal has subsequently become blocked with sand and is no longer kept open (DWAF, 2006). The state of information on the Seekoei Estuary is considered good (Whitfield, 2000). Physico-chemical Physico-chemical parameters were measured at four locations in the Seekoei Estuary from the mouth to the upper reaches of the system. The mouth of the estuary was closed at the time of sampling. According to anecdotal information the mouth of the Seekoei was historically predominately open before major anthropogenic disturbance. However, the deflection of the estuary mouth and the construction of the causeway and concrete canal caused the mouth to close for extended periods and to stay open for only a few days (Bickerton & Pierce, 1988). More recently, since this survey was undertaken, the increase in the opening in the causeway and the cessation of artificially maintaining the mouth via the concrete canal has allowed the estuary to remain open for much longer periods of up to 18 months even when river flow is very low (DWAF, 2006). The channel water depth during this survey ranged between 0.5 and 1.6 m (Table 1). Bickerton & Pierce (1988) also reported depths of between 0.8 and 1.6 m in November 1984. Both surface and bottom temperatures measured between 17.6 and 20.8°C and generally increased from the mouth upstream (Table 1). Surface temperatures were only slightly warmer than those at the bottom at the uppermost site (Site 4). Similar conditions were also reported in spring (November 1984) where water temperatures increased in an upstream direction and ranged between 17.5 and 18.6°C (Bickerton & Pierce, 1988); similarly, there was little difference recorded between surface and bottom waters. Both surface and bottom salinities were fairly uniform and decreased from 9.3‰ recorded at the mouth to 3.9‰ at the uppermost site (Table 1). Salinities reported by Bickerton & Pierce (1988) were relatively constant throughout the estuary, ranging between 26.0 and 27.0‰. Salinity can be extremely variable in the Seekoei Estuary, with severe hypersaline conditions sometimes developing in the upper reaches (DWAF, 2006). Dundas (1994) measured salinities of over 45.0‰ in summer (October–March 1992) and this was attributed to high evaporation rates. The pH of the water ranged between 8.2 and 9.4 and did not vary much between surface and bottom waters (Table 1). Bickerton & Pierce (1988) reported most sites having a pH typical of seawater (8.0) with the exception of the uppermost site (6.8) and this was attributed to a more fluvial influence. Dissolved oxygen concentrations ranged between 8.0 and 12.8 mg l–1 (Table 1). Dissolved oxygen concentrations of between 7.4 and 7.9 mg l–1 were recorded in November 1984, with bottom and surface measurements being similar and this was attributed to wind-induced mixing (Bickerton & Pierce, 1988). The waters of the Seekoei were relatively clear with the bed of the system visible at the lower sites (Sites 1 & 2); Secchi disc measurements for the remaining sites measured between 0.6 and 0.9 m (Table 1). Bickerton & Pierce (1988) also found that the Secchi disc readings in the system were mostly above 0.7 m. Ichthyofauna Eleven seine net hauls and five gill nets caught a total of 1192 individuals representing 13 species from nine families. Mugilidae (3 species), Sparidae (2 species) and Gobiidae (2 species) dominated the taxa. Catches were numerically dominated by 76 Transactions of the Royal Society of South Africa G. aestuaria, which comprised 28.9% of the catch, followed by A. breviceps (21.8%), juvenile mugilids (21.7%), L. lithognathus (16.5%), M. cephalus (3.1%), G. callidus (2.6%) and L. dumerili (1.7%) (Table 2). A total species mass of over 27 kg was caught and this was dominated by M. cephalus (51.1%), L. lithognathus (13.6%), L. richardsonii (10.2%), L. dumerili (9.2%), Oreochromis mossambicus (7.4%), L. amia (5.1%) and R. holubi (1.2%) (Table 3). Based on the results of a gill net survey (Bickerton & Pierce, 1988) and a study using seine and gill nets (Dundas, 1994), a total of 24 species have been recorded in the Seekoei Estuary. Thirteen of these species (54%) were captured during this survey. Important species, in terms of abundance and/or biomass included A. breviceps, G. feliceps, G. aestuaria, L. amia, L. lithognathus, L. dumerili, L. richardsonii, M. falciformis, M. cephalus and R. holubi (Bickerton & Pierce, 1988; Dundas, 1994). Four estuarine-resident species (A. breviceps, G. aestuaria, G. callidus and P. knysnaensis) were captured during this survey. Atherina breviceps ranged from 27 to 57 mm in size; the predominant size classes included both immature (30–40 mm) and mature (40–50 mm) individuals. Gilchristella aestuaria measured between 26 and 40 mm; most specimens were mature individuals in the 30–50 mm size classes; G. aestuaria are estimated to mature at a size of 28 mm SL (Talbot, 1982). Glossogobius callidus were between 20 and 62 mm; this species matures at approximately 35 mm (Whitfield, 1998) and most specimens during this survey were mature individuals in the 30–50 mm size classes. The presence of both mature and immature individuals of a range of estuarine-resident species suggests that the Seekoei serves as a viable habitat for this group of fishes. Eight estuarine-associated marine species were captured; these were represented by H. capensis, L. amia, L. lithognathus, L. dumerili, L. richardsonii, M. falciformis, M. cephalus and R. holubi. Lithognathus lithognathus ranged in size from 19 to 289 mm SL; most specimens were newly recruited individuals (<40 mm) although some larger specimens (>240 mm) were also present. This indicates regular recruitment and utilisation of the system. Liza dumerili were all large (>180 mm) individuals and although no juvenile specimens were identified, the presence of high numbers of juvenile mugilids (mostly <50 mm) suggests that the Seekoei serves a viable nursery function for this group of fishes. Mugil cephalus ranged in size from 22 to 360 mm. Most specimens were large (>200 mm), but a few small (<40 mm), newly recruited individuals were also recorded. Recruitment of M. cephalus into estuaries occurs at <30 mm TL (Wallace & van der Elst, 1975). Overall, both estuarine-resident species and estuarineassociated marine species dominated the ichthyofauna of the Seekoei during this survey. Estuarine-resident species accounted for 31% of the taxa captured while estuarineassociated marine species comprised 62% of the taxa; estuarine resident species comprised 54.1% of the abundance and 1.7% of the biomass, while estuarine-associated marine species comprised 45.5% of the abundance and 91.0% of the mass. These results indicate that despite major anthropogenic disturbance, the Seekoei Estuary still serves an important nursery function for estuarine-associated fishes. Kabeljous The Kabeljous (34°00’S, 24°56’E) is situated just north of Jeffrey’s Bay. The river is 30 km long and has a catchment area of 238 km2 (Reddering & Esterhuysen, 1984). Several tributaries flow into the Kabeljous and there are a number of farm dams in the catchment (Bickerton & Pierce, 1988). The upper reaches of the system are narrow but at the coast the system forms a wide, shallow expanse of water. The system is relatively undisturbed; Vol. 65(1): 69–84, 2010 the holiday township of Kabeljous-on-Sea is situated at the coast on the lower southwestern side of the estuary but there is little to no development in the middle and upper reaches of the estuary (Dundas, 1994). Two road bridges cross the Kabeljous between 2 and 3 km upstream of the mouth (Bickerton & Pierce, 1988). According to Whitfield (2000) the state of information on the Kabeljous is regarded as moderate. Physico-chemical Physico-chemical parameters were measured at three locations in the lower, middle and upper reaches of the estuary. The mouth of the system was closed at the time of sampling. The Kabeljous is typically separated from the sea by a large, 100–200 m wide sandbar. According to Bickerton & Pierce (1988) the estuary remains closed for most of the year and normally only opens during floods. The water depth during this survey ranged between 0.9 and 1.1 m (Table 1). The average depth of the system has been reported to be 0.5 m, while depths of between 1.6 and 2.3 m were recorded in the upstream channel (Bickerton & Pierce, 1988). Surface water temperatures increased from 17.2°C near the mouth to 17.9°C at the uppermost site; bottom water temperatures were slightly higher and increased from 17.3°C to 18.4°C in the same direction (Table 1). Surface and bottom water temperatures measured in November 1984 ranged between 18.5°C and 23.0°C and also generally increased upstream from the mouth (Bickerton & Pierce, 1988). Salinities measured during this survey decreased upstream from the mouth; surface salinities ranged from 17.1 to 15.2‰ and bottom salinities from 16.9 to 16.5‰ (Table 1). Bickerton & Pierce (1988) reported no differences between surface and bottom salinities, which ranged between 30.0 and 32.0‰ in November 1984. The pH of the water during this survey was fairly uniform throughout the estuary and measured between 7.9 and 8.1 (Table 1). The pH values recorded by Bickerton & Pierce (1988) in November 1984 were slightly higher and ranged between 8.4 and 8.6; this is probably a reflection of the higher salinities recorded at the time. The dissolved oxygen concentrations of the surface water during this survey ranged between 4.6 and 7.0 mg l–1 and were slightly higher than those at the bottom, which measured between 4.1 and 5.7 mg l–1 (Table 1). This is probably a result of a lack of mixing and thermal stratification of the water column. Dissolved oxygen values reported by Bickerton & Pierce (1988) in November 1984 were higher than those recorded during this study and measured between 8.3 and 10.5 mg l–1. The waters in the Kabeljous were clear and the bed of the system was visible at all sites (Table 1). Bickerton & Pierce (1988) also recorded high Secchi disc measurements (>1.6 m) during their survey of the Kabeljous. Ichthyofauna A total of 13 seine net hauls and five gill nets caught 2248 individuals representing 11 species from seven families. Mugilidae (5 species) and Sparidae (2 species) dominated the taxa. Catches were numerically dominated by A. breviceps, which comprised 68.0% of the total abundance; this was followed by juvenile mugilids (11.6%), L. richardsonii (11.3%), L. lithognathus (5.3%), R. holubi (1.6%) and M. cephalus (1.1%) (Table 2). A total species mass of over 46 kg was caught and this was dominated by L. richardsonii (65.1%), M. cephalus (11.2%), L. amia (10.8%), M. capensis (5.0%), L. lithognathus (3.2%) and A. breviceps (3.0%) (Table 3). Based on information provided from a gill net and beam trawl survey (Bickerton & Pierce, 1988) and the results of a seine net and gill net study (Dundas, 1994), a total of 23 species James & Harrison: Survey of the estuaries on southeast coast of South Africa, Cape St Francis – Cape Padrone have been recorded in the Seekoei Estuary. Eleven of these species (48%) were also recorded during this survey. The dominant species either numerically and/or in terms of biomass included A. japonicus, A. breviceps, G. aestuaria, L. amia, L. lithognathus, L. richardsonii, L. tricuspidens, M. falciformis, M. cephalus, O. mossambicus, P. commersonnii, Pomadasys olivaceum and R. holubi (Bickerton & Pierce, 1988; Dundas, 1994). Of the taxa collected during this survey, three species (27%) were estuarine-resident species and these included A. breviceps, G. aestuaria and P. knysnaensis. Atherina breviceps comprised both immature and mature individuals between 26 and 68 mm, indicating that the Kabeljous serves an important habitat for estuarine-resident species. Eight species (73%) were estuarineassociated marine species and these included L. amia, L. lithognathus, L. richardsonii, L. tricuspidens, M. falciformis, M. cephalus, M. capensis and R. holubi. Lithognathus lithognathus comprised mostly small individuals in the 20–40 mm size classes, with a few large individuals (>250 mm) also present. Liza richardsonii comprised individuals between 40 and 325 mm. Small individuals were mostly in the 40–50 mm size class and large individuals in the 230–240 mm size class, indicating regular use of the system. Rhabdosargus holubi were mostly newly recruited individuals in the 15–25 mm size classes. Similarly, M. cephalus comprised small individuals in the 20–40 mm size classes, with a few individuals larger than 250 mm. The high proportion of newly recruited fish, together with the occurrence of a few larger specimens indicates that the Kabeljous provides important nursery habitat for estuarine-associated marine species. Overall, estuarine species comprised 68.2% of the catch numerically and 3.0% by mass during this survey; estuarineassociated marine species comprised 31.8% of the catch numerically and 97.0% by mass. The high proportion of both marine and estuarine-resident species indicates that the Kabeljous serves an important nursery function for these fishes. Gamtoos The Gamtoos (33°58’S, 25°04’E) is located approximately 60 km west of Port Elizabeth. The system has a river length of 75 km and a catchment area of 34 438 km2 (Heinecken, 1981). Most of the catchment area is situated in a region of low precipitation and there are three major dams within the catchment that absorb and buffer a large proportion of the runoff. Despite this, occasional floods completely flush the system with freshwater (Schlacher & Wooldridge, 1996). The upper reaches of the estuary are narrow and flow through steep-sided banks; the system gradually broadens from the middle reaches towards the mouth and an extensive flood-tidal delta is situated in the lower reaches (Heinecken, 1981). Three major structures cross the system, the old National Road Bridge is located about 7 km from the mouth, a water pipeline and the N2 National Road cross the system approximately 5 km from the mouth (Heinecken, 1981). The state of information on the Gamtoos is regarded as good (Whitfield, 2000). Physico-chemical Physico-chemical parameters were measured at eight locations in the Gamtoos. Site 1 was located in a tidal side arm that represents the remains of the former main channel, Site 2 in the estuary mouth and Sites 3–8 in the lower, middle and upper reaches. The mouth of the Gamtoos is permanently open and is maintained by tidal currents rather than river flow; this is supported by the presence of a flood-tidal delta in the lower reaches (Heinecken, 1981). The channel depth during this survey ranged between 1.0 and 2.6 m (Table 1). Heinecken 77 (1981) reported depths of between 2.0 and 4.0 m in the estuary channel in August 1980. According to Schlacher & Wooldridge (1996) the tidal inlet is shallow (<1.5 m); beyond the flood tidal delta the channel deepens to about 4.0 m in the middle reaches and then becomes progressively shallower toward the tidal head (approximately 20 km from the mouth) where the water depth is less than 1.0 m. Surface water temperatures during this survey ranged between 18.5 and 20.2°C, and were generally higher than those at the bottom, which measured between 18.0 and 19.4°C (Table 1). Water temperatures tended to increase upstream from the mouth. In August 1980, surface water temperatures measured between 17.3 and 17.9°C and also increased in an upstream direction (Heinecken, 1981). Schlacher & Wooldridge (1996) found that winter temperatures decreased from 16.6 to 12.2°C in an upstream direction, whereas in summer this trend was reversed and temperatures increased from 19.1 to 24.8°C in an upstream direction. Both surface and bottom salinities during this survey declined from the mouth upstream and ranged from 31.4 to 3.1‰ (Table 1); surface values were also generally lower than those recorded near the bottom. Schlacher & Wooldridge (1996) found that axial salinity gradients in the Gamtoos can vary dramatically with the magnitude of river inflow. Marais (1983b) recorded bottom salinities of only 2.0‰ at the mouth when the river was in flood in October 1981; by contrast, bottom salinity at the mouth averaged 30.0‰ when the river was not in flood. The pH of the surface waters during this survey measured between 7.8 and 8.4, while the pH of the bottom waters ranged between 7.6 and 8.2 (Table 1). Surface dissolved oxygen concentrations ranged from 7.6 to 11.6 mg l–1 and were higher on the surface than at the bottom; bottom dissolved oxygen concentrations ranged between 3.3 and 9.7 mg l–1 (Table 1). Secchi disc measurements during this study were less than 1.0 m (Table 1). Heinecken (1981) also reported low Secchi disc readings of 0.8 m throughout the estuary. Ichthyofauna A total of 24 species from 14 families were caught in 16 seine hauls and 15 gill nets; 4272 individuals were recorded with Mugilidae (5 species), Sparidae (4 species) and Gobiidae (3 species) dominating the taxa. Numerically, catches were dominated by G. aestuaria (26.6%) followed by L. dumerili (16.2%), R. holubi (15.3%), L. richardsonii (12.1%), juvenile mugilids (8.0%), A. breviceps (5.1%), M. cephalus (3.3%), G. feliceps (2.6%), L. lithognathus (2.5%), R. globiceps (1.8%), P. knysnaensis (1.2%) and P. commersonii (1.0%) (Table 1). A total species mass of over 170 kg was recorded with G. feliceps dominating; this species comprised 36.9% of the total biomass followed by A. japonicus (17.9%), L. richardsonii (11.4%), Elops machnata (8.1%), L. tricuspidens (7.0%), L. dumerili (5.6%), R. holubi (3.6%), M. cephalus (2.4%), L. lithognathus (1.8%), P. commersonnii (1.5%) and M. capensis (1.5%) (Table 3). Marais (1983b) recorded 22 species from a gill net survey of the estuary, 14 of which (64%) were recorded during this survey. Similar species were also among the dominant taxa, both numerically and in terms of biomass, and these included A. japonicus, G. feliceps, L. amia and Mugilidae (L. dumerili, L. richardsonii, L. tricuspidens, M. cephalus and M. capensis) (Marais, 1983b). From this survey, five species (20%) live and breed in estuaries; these are represented by A. breviceps, C. gilchristi, G. aestuaria, G. callidus and P. knysnaensis. Atherina breviceps ranged between 28 and 59 mm and included both mature and immature individuals; most specimens fell within the 30–40 mm size class. 78 Transactions of the Royal Society of South Africa Gilchristella aestuaria measured 19–60 mm and the majority of individuals were mature (>30 mm). All P. knysnaensis individuals were mature and measured between 30 and 52 mm. The presence of both juvenile and adult estuarine-resident species indicates that the Gamtoos Estuary is an important habitat for these fishes. Nineteen estuarine-associated marine species were captured during this survey. These included A. japonicus, D. capensis, E. machnata, G. feliceps, H. capensis, L. amia, L. lithognathus, L. dumerili, L. richardsonii, L. tricuspidens, M. falciformis, M. cephalus, M. capensis, P. indicus, P. commersonnii, P. saltatrix, R. globiceps, R. holubi and S. turbynei. The majority of G. feliceps individuals during this survey were between 300 and 310 mm. This species has a unique reproductive cycle in that the male mouthbroods the eggs; mature males (>260 mm SL) with eggs frequent estuaries during spring and summer (Whitfield, 1998). All G. feliceps specimens captured were mouthbrooding males, which indicates that the Gamtoos is an important nursery for this species. Two size classes of L. lithognathus were caught, the first was between 20 and 30 mm and the second between 130 and 140 mm; this indicates regular recruitment and utilisation of the system. Liza dumerili ranged in size from 31–245 mm; most specimens were small individuals in the 60–70 mm size class, which suggests recent recruitment. Two size classes of L. richardsonii were caught; the first comprised individuals of 80–90 mm and the second individuals >200 mm. This also indicates regular utilisation of the estuary. Most R. holubi captured were small, recently recruited individuals between 40 and 50 mm. The presence of both small, recently recruited individuals and larger specimens of estuarineassociated marine species indicates that the Gamtoos is an important nursery for this group of fishes. Both estuarine-resident and estuarine-associated marine species dominated the fish fauna of the Gamtoos Estuary. Estuarine-associated marine species comprised 66.2% of the total catch numerically and 99.4% by mass, while estuarineresident species comprised 33.9% of the catch numerically and 0.6% by mass. These results indicate that the Gamtoos Estuary serves an important habitat for both resident and estuarineassociated marine species. Van Stadens The Van Stadens (33°58’S, 25°13’E) is situated about 32 km west of Port Elizabeth. The river is 25.8 km long with a catchment area of 271 km2 (Dundas, 1994). The estuary flows between relatively steep banks for most of its length and shallow littoral areas are limited. The Van Stadens forms part of a holiday resort and a caravan park is situated near the coast on the eastern shore of the system. All development, however, is set away from the banks of the system, which are relatively undisturbed (Dundas, 1994). A water pipeline crosses the system about 500 m from the mouth and the N2 National Road crosses the river approximately 11 km from the mouth. The state of information on the Van Stadens is regarded as poor with limited biological information (Whitfield, 2000). Physico-chemical Physico-chemical parameters were measured at four sites in the Van Stadens from the mouth to the upper reaches. The mouth of the system was closed at the time of sampling. The estuary mouth is separated from the sea by a 50 m wide beach with a low berm. Freshwater inflow can result in the sand barrier being breached although high seas overwashing and lowering the sand bar can also cause the mouth to open (Dundas, 1994). The water depth during this survey ranged Vol. 65(1): 69–84, 2010 between 1.5 and 3.3 m, with the deepest section in the lower reaches; water depths at Sites 1 and 2 exceeded 2.5 m (Table 1). Surface water temperatures increased from 19.5°C at the mouth to 22.0°C in the upper reaches. Bottom water temperatures were similar, increasing from 18.5°C to 21.6°C in the same direction (Table 1). Dundas (1994) recorded a mean surface temperature of 24.0°C in summer (1992), with temperatures also increasing from the lower to the upper reaches. Surface salinities ranged between 10.5 and 15.2‰ with the lowest value recorded at the uppermost site (Site 4); the salinity of the bottom waters was fairly uniform and ranged between 15.1 and 15.2‰ (Table 1). Dundas (1994) recorded a mean surface salinity of 21.1‰ in summer, with salinity decreasing from the lower to the upper reaches. The pH of the waters during this survey decreased from the mouth to the upper reaches and ranged from 7.8 to 7.2 (Table 1). Dundas (1994) recorded a mean pH of 8.5 in summer and found that values were fairly uniform throughout the system. Both surface and bottom dissolved oxygen values decreased upstream from the mouth. The dissolved oxygen values of the surface waters ranged from 7.5 to 5.3 mg l–1 and were higher than those at the bottom, which ranged from 7.2 to 3.1 mg l–1 (Table 1). The steep banks that characterise the Van Stadens Estuary probably limit wind-induced mixing of the waters resulting in lower bottom dissolved oxygen values. Dundas (1994) recorded a mean dissolved oxygen concentration of 8.5 mg l–1 in summer. The waters in the estuary were clear and the bed of the system was visible at most sites; a Secchi disc value of 2.5 m was recorded at Site 2 (Table 1). Dundas (1994) also reported relatively high water transparencies in the Van Stadens, with mean Secchi disc values varying between 0.9 and 2.5 m. Ichthyofauna A total of 1718 individuals representing 15 species and seven families were caught in 11 seine net hauls and five gill nets. Mugilidae (5 species), Sparidae (4 species) and Gobiidae (2 species) dominated the taxa. Juvenile mugilids (53.7%) dominated catches by number followed by R. holubi (14.1%), L. lithognathus (11.5%), A. breviceps (8.2%), M. capensis (4.6%), L. richardsonii (2.8%), M. cephalus (2.2%) and G. callidus (1.0%). A total species mass of over 33 kilograms was recorded and this was dominated by L. richardsonii, which comprised over 70.2% of the catch, followed by L. amia (12.2%), L. tricuspidens (10.3%) and juvenile mugilids (4.5%) (Table 3). Dundas (1994) recorded fifteen fish species from the estuary in a seine and gill net study. Ten species (67%) were common to those captured during this survey. The dominant species captured by Dundas (1994) included A. breviceps, L. amia, L. lithognathus, L. dumerili, L. richardsonii, M. falciformis, M. cephalus, Myliobatis aquila, P. commersonnii and R. holubi. Of the fishes recorded during this survey, four (A. breviceps, G. aestuaria, G. callidus and P. knysnaensis) are estuarine-resident species. Atherina breviceps comprised mostly mature individuals between 39 and 62 mm. Eleven species (D. capensis, L. amia, L. lithognathus, L. dumerili, L. richardsonii, L. tricuspidens, M. falciformis, M. cephalus, M. capensis, R. holubi and S. salpa) are estuarine-associated marine species. Lithognathus lithognathus comprised mostly small individuals less than 30 mm; a few larger individuals (>120 mm) were also present, which suggests recent recruitment and regular utilisation of the system. Liza richardsonii ranged in size between 202 and 339 mm. Although most of these were large specimens in the 290–300 mm size class the high numbers of juvenile mugilids (<50 mm) indicates that the Van Stadens serves as a nursery for this group of fishes. This is also supported by catches of M. capensis being James & Harrison: Survey of the estuaries on southeast coast of South Africa, Cape St Francis – Cape Padrone dominated by newly recruited juveniles between 20 and 56 mm and M. cephalus comprising juveniles mostly <30 mm. Rhabdosargus holubi were also all juveniles between 15 and 72 mm in length. The presence of large numbers of juvenile estuarine-associated marine species indicates that the Van Stadens is an important nursery habitat for these fishes. Estuarine-associated marine species also dominated catches comprising 91% of the catch numerically and 99% of the biomass. Estuarine-resident species comprised 9% of the catch numerically and only 1% of the biomass. The Van Stadens Estuary appears to serve an important nursery function for estuarine-associated marine species as well as estuarineresident species. Maitland The Maitland Estuary (33°59’S, 25°17’E) is situated about 26 km west of Port Elizabeth. The system consists of a small, very shallow sandy system located behind a broad dissipative beach and little published information exists on the estuary (Whitfield, 2000). Physico-chemical Physico-chemical measurements were made at one site in the Maitland Estuary. This was situated in the outflow channel of the mouth which was open at the time of sampling. The Maitland appears to be closed to the sea for extended periods and, when the system opens, the mouth comprises a shallow drainage channel for outflowing estuarine water. The water depth measured 0.3 m; the water temperature was 22.7°C and the salinity was 1.3‰, indicating very little seawater input. The water had a pH of 8.4 and the dissolved oxygen measured 12.9 mg l–1; the bed of the system was also visible (Table 1). Ichthyofauna A total of five seine hauls caught 585 individuals representing six species and three families, with Mugilidae dominating the taxa (3 species). In terms of numbers, M. cephalus comprised 88.2% of the catch followed by L. lithognathus (6.8%), M. capensis (2.7%) and R. holubi (1.4%) (Table 2). A total species mass of 0.4 kg was caught; M. cephalus (95.3%) dominated the biomass followed by L. lithognathus (2.3%) and M. capensis (1.0%) (Table 3). From this survey, five species (L. lithognathus, L. richardsonii, M. cephalus, M. capensis and R. holubi) are estuarine-associated marine species and only one estuarine-resident species (P. knysnaensis) was recorded. All estuarine-associated species caught comprised newly recruited individuals. L. lithognathus ranged in size from 14–28 mm, M. cephalus measured 21–46 mm, Myxus capensis ranged between 20 and 27 mm, and R. holubi were between 14 and 17 mm. Although the predominance of small individuals indicates recent recruitment, the lack of larger specimens suggests that the system is not regularly utilised and that the nursery function of this system may be limited. It may also be possible; however, that larger specimens had migrated back to sea immediately after the mouth had opened and thus were not captured at the time of this survey. Estuarine-associated marine species dominated catches comprising 99.8% of the catch numerically and 99.6% of the biomass. Estuarine-resident species comprised only 0.2% of the catch numerically and 0.4% of the biomass. Although the Maitland is a small system that is intermittently open to the sea, it still appears to support estuarine-associated species. Bakens The Bakens Estuary (33°57’S, 25°37’E) is a heavily modified system that discharges into the Port Elizabeth Harbour. Harbour 79 and urban developments have resulted in the system being canalised along much of its length and the bed is composed mainly of rock and concrete. Nine bridges cross the system. The state of information on the Bakens is regarded as poor (Whitfield, 2000). Physico-chemical Physico-chemical parameters were measured at three sites in the Bakens. Only the lower site (Site 1) was deep enough to permit both surface and bottom measurements. Water depths ranged from 0.2 to 1.0 m (Table 1). Water temperatures measured between 19.0 and 20.5°C and were highest at the uppermost site (Site 3). Salinities ranged from 31.7‰ at the mouth to 0.7‰ at the uppermost site. The pH values measured between 7.8 and 8.1. Dissolved oxygen concentrations ranged from 6.2 to 15.9 mg l–1 and were highest at the uppermost site. The bed of the system was visible at all sites (Table 1). Ichthyofauna Two seine net hauls caught a total of 83 specimens representing nine species and five families. Mugilidae (4 species) and Sparidae (3 species) dominated the taxa. The most abundant species recorded was H. capensis and comprised 51.2% of the catch followed by P. knysnaensis (13.1%), L. lithognathus (9.5%), A. breviceps (4.8%), M. cephalus (4.8%), juvenile mugilids (4.8%), L. dumerili (3.6%), R. holubi (3.6%), L. richardsonii (3.6%) and D. capensis (1.2%) (Table 2). A total species mass of over 159 g was caught; this was dominated by L. richardsonii (56.0%), L. dumerili (13.1%), M. cephalus (8.8%), R. holubi (7.7%), H. capensis (5.8%), P. knysnaensis (4.3%), A. breviceps (2.4%) and L. lithognathus (1.2%) (Table 3). Of the taxa captured during this survey, seven were estuarine-associated marine species; these included D. capensis, H. capensis, L. lithognathus, L. dumerili, L. richardsonii, M. cephalus and R. holubi. Heteromycteris capensis were all immature individuals and were mostly in the 30–40 mm size class; this species attains sexual maturity at approximately 80 mm SL (Cyrus & Martin, 1991). Lithognathus lithognathus comprised newly recruited individuals in the 20–30 mm size class, which suggests that the Bakens appears to provide some nursery function for marine species. This is supported by the occurrence of small individuals of M. cephalus (20–30 mm) and R. holubi (10–20 mm). Two estuarine-resident species (A. breviceps and P. knysnaensis) were captured during this survey. Psammogobius knysnaensis were all mature individuals (>30 mm) and were mostly in the 30–40 mm size class. Overall, estuarine-associated marine species were the dominant group, comprising 81.9% of the catch numerically and 91.0% by mass. Estuarine-resident species comprised 18.1% of the catch numerically and 9.0% by mass. Although this survey indicates that the Bakens supports estuarine-associated species, the canalisation of the system and lack of habitat suggests that this is limited. Papkuils The Papkuils (33°55’S, 25°36’E) is another severely modified system situated in Port Elizabeth. The system is canalised along its lower reaches and the bed is concreted; it also appears to receive industrial and domestic effluent. Numerous road bridges cross the system. The Papkuils has a permanent outlet to the sea, which is maintained by concrete walls and rock armouring. Physico-chemical parameters were measured at a single site in the lower reaches of the system. A depth of 0.1 m was recorded and the water temperature was 24.0°C. The salinity recorded was 9.7‰ indicating limited seawater input. 80 Transactions of the Royal Society of South Africa Vol. 65(1): 69–84, 2010 The pH of the water was 7.4 and dissolved oxygen was low, measuring 3.4 mg l–1. Owing to the shallow nature of the system, the bed was visible at the sampling site (Table 1). No fish sampling was conducted during this survey. Owing to the modified state of this system it is unlikely to provide suitable habitat for estuarine-associated species. mean value of 7.2 mg l–1 with the waters being slightly less oxygenated during summer. The waters of the Swartkops were relatively clear with Secchi disc values exceeding 1.0 m (Table 1). Baird et al. (1986) also found that the waters of the Swartkops Estuary were clear with mean Secchi disc measurements of between 1.0 and 1.5 m. Swartkops The Swartkops (33°51’S, 25°38’E) is situated just north of Port Elizabeth. The river is 155 km long and the catchment area is estimated at 1438 km2 (Binning & Baird, 2001). The main tributaries of the system are the Elands River and the Chatty River (Baird et al., 1986). The only major impoundment on the Swartkops River is the Groendal Dam that supplies the industrial area of Uitenhage (Binning & Baird, 2001). The storage capacity of the dam is small relative to mean annual rainfall and it has little effect on overall run-off (Baird et al., 1986). The Swartkops Estuary is 16 km long and a concrete causeway marks the upper limit of the system (Dye, 1978). The upper reaches of the system winds through steep banks of muddy sand; the estuary widens considerably in the middle reaches, traversing a broad flood plain and broadening from 80 to 200 m. Large intertidal mudflats, islands and saltmarshes increase in extent along the lower reaches. Several bridges cross the estuary, one of which carries a dual carriageway and is located 500 m from the mouth. The state of information on the Swartkops is regarded as excellent (Whitfield, 2000). Ichthyofauna A total of 4841 fish, represented by 18 families and 30 species were caught in 19 seine net hauls and 14 gill nets in the Swartkops Estuary. In terms of the number of species recorded Sparidae (5 species) and Mugilidae (4 species) were the dominant families. Gilchristella aestuaria was the most abundant species caught comprising 28.7% of the total catch, followed by R. holubi (17.0%), P. commersonnii (15.6%), juvenile mugilids (13.7%), L. richardsonii (6.0%), L. dumerili (4.9%), R. globiceps (2.6%), C. gilchristi (2.3%), L. lithognathus (1.7%), G. callidus (1.4%), P. knysnaensis (1.1%) and A. breviceps (1.1%) (Table 2). A total species mass of over 51 kg was recorded and this was dominated by E. machnata (23.3%), G. feliceps (18.1%), P. commersonnii (10.6%), L. richardsonii (10.5%), L. tricuspidens (7.5%), L. dumerili (7.2%), R. holubi (5.0%), L. amia (4.6%), A. japonicus (4.3%), juvenile mugilids (2.9%), G. aestuaria (2.5%) and M. cephalus (1.9%) (Table 3). A total of 86 species of fish has been recorded in the Swartkops Estuary (Baird et al., 1986). The high number of species recorded is probably a reflection of a combination of the number of studies undertaken and the strong marine influence within the system. Many of the dominant species, however, were also among the important species captured during this survey. Abundant fishes reported in the Swartkops estuary included A. japonicus, A. breviceps, C. gilchristi, D. capensis, E. machnata, Engraulis japonicus, G. feliceps, G. aestuaria, L. amia, L. lithognathus, L. dumerili, L. richardsonii, L. tricuspidens, M. cephalus, P. commersonnii, P. saltatrix, P. knysnaensis and R. holubi (Marais & Baird, 1980; Beckley, 1983; Baird et al., 1988). Estuarine-resident species during this survey were represented by nine species (Ambassis dussumieri, A. breviceps, C. gilchristi, C. natalensis, C. nudiceps, C. superciliosus, G. aestuaria, G. callidus and P. knysnaensis). Caffrogobius gilchristi specimens measured between 25 and 55 mm; most specimens were immature and were in the 30–40 mm size class. Caffrogobius gilchristi matures at a length of approximately 50 mm SL (Bennett, 1989). The majority of G. aestuaria, G. callidus and P. knysnaensis were mature individuals with a modal size class of 30–40 mm. These results indicate that the Swartkops is an important habitat for estuarine-resident taxa. Estuarine-associated marine species predominated during this survey represented by 21 species (A. japonicus, D. capensis, E. machnata, G. feliceps, Hemiramphus far, H. capensis, L. amia, L. lithognathus, L. dumerili, L. richardsonii, L. tricuspidens, M. falciformis, M. cephalus, P. indicus, P. commersonnii, P. saltatrix, R. globiceps, R. holubi, S. salpa, Silago sihama and S. turbynei). Lithognathus lithognathus were between 19 and 106 mm and catches were dominated by small individuals (<40 mm), which indicates recent recruitment. Liza dumerili individuals ranged from 50 to 221 mm with most also represented by recently recruited individuals in the 60–80 mm size classes. Liza richardsonii ranged between 38 and 303 mm, with most individuals in the 60–70 mm size class. Pomadasys commersonnii ranged in size from 19 to 546 mm, with most specimens between 20 and 40 mm. Recruitment of P. commersonnii individuals into estuaries occurs at 20–30 mm TL (Wallace & van der Elst, 1975). Rhabdosargus holubi individuals measured between 14 and 124 mm, with most individuals in the 20–30 mm size classes. Physico-chemical Physico-chemical parameters were measured at six sites in the Swartkops Estuary extending from the mouth area to the head of the system. The mouth of the estuary is kept permanently open by the action of strong tidal currents, which exceed the average river flow by 60 times (Baird et al., 1986). Water depth recorded during this survey ranged from 0.6 to 3.5 m (Table 1). Baird et al. (1986) recorded depths of 2.0 m in the lower reaches while the deepest areas (>3.7 m) were recorded in the upper reaches and the mouth area. Both surface and bottom water temperatures during this survey increased slightly from the mouth upstream and ranged between 18.2 and 19.2°C (Table 1). In general the head of the system is warmer than the mouth region during summer and cooler during winter (Marais & Baird 1980; Emmerson, 1985; Baird et al., 1986). Mean summer temperatures varied between 21 and 26°C while mean winter temperatures were 11–18°C (Marais & Baird, 1980; Emmerson, 1985; Scharler et al., 1997). A horizontal salinity gradient was recorded during this survey. Surface salinities decreased from 32.7‰ in the mouth region to 25.2‰ in the upper reaches. Bottom salinities decreased from 32.7‰ to 27.0‰ in the same direction (Table 1). Emmerson (1985) recorded a permanent salinity gradient from the mouth to the head of the estuary and this was attributed to a continual input of freshwater. There was no difference observed between surface and bottom salinities in the lower and middle reaches of the estuary. Some stratification, however, was recorded in the upper reaches (Sites 5 & 6). Emmerson (1985) attributed this lack of vertical stratification in the lower reaches to wind and tidal mixing. The pH values recorded during this survey ranged from 7.9 near the mouth to 7.5 in the upper reaches (Table 1). Emmerson (1985) reported a mean pH of 8.1 but found that values were generally higher at the head of the system. Dissolved oxygen concentrations during this survey ranged from 6.9 to 5.0 mg l–1 (Table 1). Baird et al. (1981) reported a relatively constant oxygen value of about 4.5 mg l–1 within the system. Both Emmerson (1985) and Scharler et al. (1997) recorded an overall James & Harrison: Survey of the estuaries on southeast coast of South Africa, Cape St Francis – Cape Padrone The majority of the individuals captured during this survey were less than one year old; again indicating recent recruitment into the system. These results show that the Swartkops Estuary provides important nursery habitat for estuarineassociated marine species. Estuarine-associated marine species also dominated the catch both numerically (64.7%) and by mass (96.8%); estuarineresident species comprised 35.3% of the catch numerically and 3.2% by mass. The predominance of both estuarine-dependent marine species and estuarine-resident species indicates that the Swartkops Estuary is an important habitat for fishes with different life-history styles. Coega The Coega system (33°47’S, 25°41’E) is situated approximately 18 km northeast of Port Elizabeth. The system is highly altered from its original state; the entire lower reaches of the system (situated below a road bridge which crosses the system about 3 km from the mouth) is utilised for salt evaporation ponds. More recently, since the time of this survey, a harbour has been developed on the coast adjacent to the estuary and the system now opens into Ngqura (Coega) Harbour. In its present state seawater is pumped into a canal that runs parallel to the former estuarine channel and then into the evaporation ponds, which occupy the estuary. The state of information on the Coega system is regarded as poor (Whitfield, 2000). Physico-chemical Physico-chemical parameters were measured at one site in the mouth area below the salt evaporation pans. The depth recorded at this site was 0.7 m and the water temperature measured 13.4°C (Table 1). The waters were hypersaline (47.2‰) and the pH measured 8.2. Dissolved oxygen values were between 7.1 and 7.2 mg l–1; the waters were clear and the estuary bottom was visible (Table 1). Ichthyofauna Five seine net hauls caught a total of 1957 specimens representing eight species and six families. Mugilidae and Gobiidae dominated the taxa with two species each. The most abundant species included A. breviceps (96.8%) and G. aestuaria (1.5%), which together comprised over 98% of the catch (Table 1). A total species mass of over 1.7 kg was recorded. Atherina breviceps (91.2%) was the dominant species by mass followed by L. richardsonii (6.1%) and G. aestuaria (1.1%) (Table 2). Of the taxa collected in the Coega four (A. breviceps, G. aestuaria, G. callidus and P. knysnaensis) were estuarine species and four (H. capensis, L. richardsonii, M. cephalus and R. holubi) were estuarine-associated marine fishes. Atherina breviceps captured during this survey measured between 12 and 62 mm; although most individuals fell within the 30–50 mm size classes, a cohort of small specimens 10–20 mm were also present. Most G. aestuaria specimens were mature individuals in the 30–50 mm size classes. The presence of both immature and mature specimens suggests that the Coega is able to support estuarine-resident fishes. Estuarine-resident species were also the dominant group of fishes both numerically and by mass, comprising 98.7% and 93.5% of the catch, respectively. Estuarine-associated marine species comprised 1.3% of the catch numerically and 6.5% of the mass. This survey indicates that although the Coega is severely modified, it still supports estuarine-resident taxa. However its suitability as a nursery habitat for estuarineassociated marine taxa is limited. 81 Sundays The Sundays (33°43’S, 25°51’E) flows into Algoa Bay approximately 35 km northeast of Port Elizabeth. The river, which is 310 km long, has a catchment area of 20 729 km2 (Harrison & Whitfield, 1990). One major impoundment, the Mentz Dam, is situated on the river but this has not significantly affected the flow of freshwater into the system (Marais, 1981). The estuary is about 21 km long and is channel-like along its entire length with steep banks and an absence of extensive mudflats or salt marshes (Wooldridge & Melville-Smith, 1979; Beckley, 1984). A sandy flood-tidal delta is situated in the lower reaches and these sand banks are exposed at low tide. The estuary is widest just inside the mouth (maximum width of 800 m) from where it becomes progressively narrower to about 20 m at the head of the estuary (Whitfield & Harrison, 1996). The state of information on the Sundays Estuary is considered excellent (Whitfield, 2000). Physico-chemical Physico-chemical parameters were measured at seven sites in the Sundays Estuary ranging from the mouth area (Site 1) to the upper reaches of the system (Site 7). The mouth of the Sundays Estuary is permanently open and this is maintained by tidal currents. Water depths recorded during this survey ranged from 1.2 to 3.5 m (Table 1). According to Marais (1981) the estuary is 2.5 m deep near the mouth and gradually increases to a maximum depth of 5.0 m in the lower reaches from where it gets gradually shallower to approximately 2.5 m in the upper reaches. Both surface and bottom water temperatures during this survey increased from the mouth upstream; surface water temperatures were also slightly higher than those at the bottom. Surface water temperatures ranged from 19.1 to 21.9°C, while bottom water temperatures measured 18.9–20.4°C (Table 1). Jerling & Wooldridge (1991) recorded water temperatures ranging from 13.0°C in winter (July) to 26.0°C in summer (January), with differences between surface and bottom water temperatures rarely exceeding 1.0°C. Both horizontal and vertical salinity gradients were recorded during this survey. Surface salinities were generally lower than those at the bottom and decreased from 32.8‰ in the mouth region to 3.4‰ in the upper reaches. Bottom salinities decreased from 32.8‰ to 3.2‰ in the same direction (Table 1). Jerling & Wooldridge (1991) also noted that horizontal and vertical salinity gradients were evident throughout the year, with salinity in the mouth region consistently above 30.0‰ and seldom above 5.0‰ at the head of the estuary; differences between surface and bottom salinities increased up the estuary. The pH of both surface and bottom waters generally increased upstream from the mouth; surface values (8.0–8.5) were also slightly higher than those recorded at the bottom (7.8–8.5) (Table 1). Surface dissolved oxygen concentrations were typically higher than those recorded at the bottom and are probably a reflection of stratification and a lack of mixing. Surface values ranged from 7.8 to 9.9 mg l–1 while those at the bottom measured between 4.9 and 8.9 mg l–1 (Table 1). The waters of the Sundays were found to be well oxygenated with mean dissolved oxygen values exceeding 7.0 mg l–1 (Emmerson, 1989; Scharler et al., 1997). The water clarity during this survey generally decreasing upstream from the mouth with Secchi disc values measuring between 0.4 and 0.7 m (Table 1). Similar values were reported by other workers who reported mean Secchi disc values in the Sundays Estuary of between 0.3 and 1.0 m (Marais, 1984; Scharler et al., 1997). 82 Transactions of the Royal Society of South Africa Ichthyofauna Sixteen seine net hauls and 14 gill nets caught 27 species representing 15 families. Mugilidae and Sparidae were each represented by five species. A total of 3550 individuals was recorded and these were dominated by G. aestuaria (32.7%), juvenile mugilids (17.4%), R. holubi (14.3%), L. richardsonii (6.5%), L. dumerili (5.6%), M. cephalus (4.5%), P. commersonnii (4.0%), C. gilchristi (2.7%), L. lithognathus (2.6%), S. turbynei (1.9%), A. breviceps (1.6%), R. globiceps (1.4%) and L. tricuspidens (1.0%) (Table 2). A species mass of over 134 kg was recorded with E. machnata (34.9%), L. tricuspidens (24.8%), A. japonicus (7.3%), M. cephalus (7.0%), L. richardsonii (6.0%), Carcharias taurus (5.9%), P. commersonnii (4.0%), G. feliceps (3.0%), L. dumerili (2.5%) and R. holubi (2.0%) dominating the catch (Table 3). Based of the results of a gill net study (Marais, 1981) and a seine net study (Beckley, 1984), a total of 55 species have been recorded in the Sundays Estuary. Altogether 24 of these species (44%) were common to this survey, many of which were dominant during all studies. Important species, either numerically and/or in terms of biomass included A. japonicus, C. gilchristi, G. feliceps, G. aestuaria, H. capensis, L. dumerili, L. richardsonii, M. falciformis, M. cephalus and P. commersonnii, P. knysnaensis, R. holubi and S. bleekeri (Marais, 1981; Beckley, 1984). Of the taxa collected during this survey seven were estuarineresident species (A. breviceps, C. gilchristi, C. natalensis, G. aestuaria, G. callidus, P. knysnaensis and S. temminckii). Atherina breviceps were all mature individuals between 48 and 65 mm, with a modal size class of 50–60 mm. Caffrogobius gilchristi specimens were between 17 and 62 mm; most were immature and were in the 30–40 mm size class. The majority of G. aestuaria caught were mature individuals with a modal size class of 40–50 mm. The presence of mature and immature individuals indicates that the Sundays is an important habitat for resident taxa. Seventeen species during this survey were estuarine-associated marine species (A. japonicus, D. capensis, E. machnata, G. feliceps, H. capensis, L. lithognathus, L. dumerili, L. richardsonii, L. tricuspidens, M. falciformis, M. cephalus, M. capensis, P. indicus, P. commersonnii, R. globiceps, R. holubi and S. turbynei). Liza dumerili ranged in size from 30 to 240 mm; most of these were small individuals in the 50–60 mm size class. The majority of L. richardsonii were also small individuals between 80 and 90 mm; a few individuals >200 mm were also captured indicating regular usage of the system. Mugil cephalus were predominantly small, newly recruited individuals less than 30 mm; larger specimens >290 mm were also represented. Pomadasys commersonnii showed a bimodal length frequency distribution, the majority of fish were recently recruited individuals in the 20–30 mm size class with larger specimens represented by the 270–280 mm size class. Rhabdosargus holubi individuals caught in this survey were between 15 and 130 mm; most specimens caught were 0+ juveniles between 40 and 60 mm. The predominance of small juvenile estuarine-associated marine species together with larger individuals suggests regular recruitment and utilisation of the estuary as a nursery area. Estuarine-associated marine species comprised 61.9% of the catch numerically and 93.0% by mass, estuarine-resident species comprised 38.0% numerically and 1.0% by mass. The predominance of both estuarine-associated marine and estuarine-resident taxa highlights the importance of this system as a habitat area for estuarine-associated fishes. GENERAL DISCUSSION Eleven rivers intersect the coastline between Cape St Francis and Cape Padrone (Figure 1). Four systems (Kromme, Vol. 65(1): 69–84, 2010 Gamtoos, Swartkops and Sundays) are large, permanently open estuaries. A connection with the sea is maintained in these estuaries by strong tidal currents. The flow regime of the Kromme Estuary is highly altered with two large dams severely restricting freshwater input into the estuary (Scharler & Baird, 2000). The Gamtoos, although dammed, is subject to occasional floods that completely flush the system with freshwater (Schlacher & Wooldridge, 1996). The Swartkops is situated in the immediate confines of Port Elizabeth and flows through a highly urbanized and industrial area. The Sundays Estuary receives a steady flow of freshwater and is channel-like along its entire length (Wooldridge & Melville-Smith, 1979). Between 24 (Gamtoos) and 30 (Kromme and Swartkops) species were recorded in these four estuaries, with Mugilidae, Sparidae and Gobiidae dominating the taxa. Studies have shown that permanently open estuaries often have a higher species richness than estuaries that have an intermittent connection with the sea and this can be attributed to an increase in the number of marine-spawning species in permanently open estuaries (Bennett, 1989; Whitfield & Kok, 1992). Dominant estuarine-dependent marine species during this study included A. japonicus, E. machnata, G. feliceps, L. amia, L. dumerili, L. richardsonii, P. commersonnii and R. holubi. Studies in permanently open warm-temperate estuaries have also reported comparable fish assemblages (e.g. Marais, 1981, 1983b; Beckley, 1984; Baird et al., 1986). Differences in fish assemblages between permanently open and closed estuaries in the southeastern Cape have also been attributed to a greater numerical proportion of estuarineresident species in closed estuaries (Vorwerk et al., 2003). In the Gamtoos, Swartkops and Sundays estuaries, resident species comprised <40% of the catch numerically; the exception was the Kromme Estuary where resident species comprised over 66% of the catch. Strydom & Whitfield (2000) found that the Geelhoutboom tributary serves as an important source of larvae of estuarine-resident species to the Kromme Estuary. Dominant estuarine-resident taxa included A. breviceps, G. aestuaria, G. callidus and P. knysnaensis. The diversity and abundance of fish species found in estuaries is also affected by latitude. Maree et al. (2000) recorded a decrease in the number of fish species in a southwesterly direction, with a substantial decline occurring west of the permanently open Swartkops Estuary. This was attributed to the Agulhas Current moving further offshore in the Algoa Bay region and consequently tropical coastal species reaching the southern limit of their distribution. The increased occurrence of tropical species may account for the greater number of species recorded in the Sundays (27 species) than the Gamtoos Estuary (24 species). Three systems (Seekoei, Kabeljous and Van Stadens) are moderately sized estuaries characterised by intermittently open mouths separated from the sea by a broad dissipative beach. The Seekoei Estuary has undergone severe human disturbance in the form of reduced river flow and completely transformed mouth dynamics. In comparison the Kabeljous and Van Stadens estuaries are relatively undisturbed. The Maitland is a small, shallow sandy system that is typically closed to the sea for extended periods. Between six (Maitland) and 15 (Van Stadens) species were recorded in these four estuaries. Estuarine-resident taxa dominated the catches numerically in the Seekoei and Kabeljous estuaries, comprising >50% of the catch, while estuarineassociated marine species dominated catches numerically and by mass in the Van Stadens and Maitland Estuaries. Dominant estuarine-associated marine species included mugilids James & Harrison: Survey of the estuaries on southeast coast of South Africa, Cape St Francis – Cape Padrone (M. cephalus, L. richardsonii and M. capensis) and sparids (R. holubi and L. lithognathus). Dominant estuarine-resident species included A. breviceps, G. callidus, G. aestuaria and P. knysnaensis. Only one freshwater species, Oreochromis mossambicus was recorded in the Seekoei Estuary. The remaining three systems (Bakens, Papkuils and Coega) are all highly altered systems that provide little habitat for either resident or migrant marine species. The coastal sector between Cape St Francis and the Sundays Estuary contains eleven coastal outlets of which four are large permanently open estuaries. The greatest diversity of species was found in the permanently open estuaries, while four temporarily open/closed estuaries, although supporting lower species diversity also provided important habitat for a number of estuarine-associated marine species and resident taxa. Three of the outlets are highly altered and provide little habitat for fishes. Of the systems assessed by Whitfield (2000) within this coastal sector, the state of information on 45% of the estuaries was regarded as poor and the remaining estuaries were regarded as having moderate (9%), good (18%) to excellent (27%) information. As a result of this survey basic physicochemical and biological (fishes) information now exists on all of these systems, with much of this information having been lacking for the smaller altered systems. REFERENCES BAIRD, D., MARAIS, J.F.K. & WOOLDRIDGE, T. 1981. The influence of a marine canal system on the ecology of the Kromme estuary, St Francis Bay. South African Journal of Zoology 16: 21–34. BAIRD, D., HANEKOM, N.M. & GRINDLEY, J.R. 1986. Estuaries of the Cape. Part II: Synopses of available information on individual systems. Report No. 23: Swartkops (CSE 3). Stellenbosch. CSIR Research Report 422: 82 pp. BAIRD, D., MARAIS, J.F.K. & MARTIN, A.P. 1988. The Swartkops Estuary. Proceedings of a symposium held on 14 and 15 September 1987 at the University of Port Elizabeth. South African National Scientific Programmes Report 156: 107 pp. BECKLEY, L.E. 1983. The ichthyofauna associated with Zostera capensis Setchell in the Swartkops estuary, South Africa. South African Journal of Zoology 18: 15–24. BECKLEY, L.E. 1984. The ichthyofauna of the Sundays estuary, South Africa, with particular reference to the juvenile marine component. Estuaries 7: 248–258. BENNETT, B.A. 1989. A comparison of the fish communities in nearby permanently open, seasonally open and normally closed estuaries in the south-western Cape, South Africa. South African Journal of Marine Science 8: 43–55. BENNETT, B.A. 1993. Aspects of the biology and life history of white steenbras Lithognathus lithognathus in southern Africa. South African Journal of Marine Science 13: 83–96. BICKERTON, I.B. & PIERCE, S.M. 1988. Estuaries of the Cape. Part II: Synopses of available information on individual systems. Report No. 33: Krom (CMS 45), Seekoei (CMS 46) and Kabeljous (CMS 47). Stellenbosch. CSIR Research Report 432: 109 pp. BINNING, K. & BAIRD, D. 2001. Survey of heavy metals in the sediments of the Swartkops River Estuary, Port Elizabeth, South Africa. Water SA 27: 461–466. CYRUS, D.P. & MARTIN, T.J. 1991. The importance of estuaries in life histories of flatfish species on the southern coast of Africa. Netherlands Journal of Sea Research 27: 255–260. DANDO, P.R. 1984. Reproduction in estuarine fish. In Potts, G.W. & Wootton, R.J. (Eds) Fish Reproduction: Strategies and Tactics. London, Academic Press. pp. 155–170 DAY, J.H. 1981. Summaries of current knowledge of 43 estuaries in southern Africa. In Day, J.H. (Ed.) Estuarine Ecology with Particular Reference to Southern Africa. Cape Town, Balkema. pp. 251–330. 83 DUNDAS, A. 1994. Comparative analysis of fish abundance and diversity in three semi-enclosed estuaries in the Eastern Cape. Unpublished MSc thesis. Port Elizabeth, University of Port Elizabeth. 80 pp. DWAF. 2006. Kromme/Seekoei Catchments Reserve Determination Study. Prepared by CSIR for CES. Report No. RDM/K90/04/CON/0705. 112 pp. DYE, A.H. 1978. Seasonal fluctuations in the vertical distribution of meiofauna in the Swartkops estuary. Zoologica Africana 13: 207–212. EMMERSON, W.D. 1985. The nutrient status of the Swartkops River estuary, eastern Cape. Water SA 11: 189–198. EMMERSON, W.D. 1989. The nutrient status of the Sundays River estuary South Africa. Water Research 23: 1059–1067. EMMERSON, W.D. & ERASMUS, T. 1987. The nutrient status of the Kromme River estuary, South Africa. Hydrobiologia 148: 87–96. HAEDRICH, R.L. & HALL, C.A.S. 1976. Fishes and Estuaries. Oceanus 19: 55–63. HANEKOM, N. & BAIRD, D. 1984. Fish community structures in Zostera and non-Zostera regions of the Kromme estuary, St Francis Bay. South African Journal of Zoology 19: 295–301. HANEKOM, N. & BAIRD, D. 1988. Distribution and variations in seasonal biomass of eelgrass Zostera capensis in the Kromme estuary, St Francis Bay, South Africa. South African Journal of Marine Science 7: 51–59. HARRISON, T.D. 1997a. A preliminary survey of coastal river systems on the South African west coast, Orange River – Groot Berg, with particular reference to the fish fauna. Transactions of the Royal Society of South Africa 52: 277–321. HARRISON, T.D. 1997b. A preliminary survey of coastal river systems on the South African west coast, Cape Columbine – Cape Point, with particular reference to the fish fauna. Transactions of the Royal Society of South Africa 52: 323–344. HARRISON, T.D. 1998. A preliminary survey of the coastal river systems of False Bay, south-west coast of South Africa, with particular reference to the fish fauna. Transactions of the Royal Society of South Africa 53: 1–31. HARRISON, T.D. 1999a. A preliminary survey of the estuaries on the south-west coast of South Africa, Cape Hangklip–Cape Agulhas, with particular reference to the fish fauna. Transactions of the Royal Society of South Africa 54: 257–283. HARRISON, T.D. 1999b. A preliminary survey of the estuaries on the south coast of South Africa, Cape Agulhas – Cape St Blaize, with particular reference to the fish fauna. Transactions of the Royal Society of South Africa 54: 285–310. HARRISON, T.D. 2001. Length–weight relationships of fishes from South African estuaries. Journal of Applied Ichthyology 17: 46–48. HARRISON, T.D. & WHITFIELD, A.K. 1990. Composition, distribution and abundance of ichthyoplankton in the Sundays River estuary. South African Journal of Zoology 25: 161–168. HECHT, T. 1973. The ecology of the Kromme estuary with special reference to Sesarma catenata. Unpublished MSc thesis. Port Elizabeth, University of Port Elizabeth. 199 pp. HEINECKEN, T.J.E. 1981. Estuaries of the Cape. Part II: Synopses of available information on individual systems. Report No. 7: Gamtoos (CMS 48). CSIR Research Report 406: 40 pp. HEYDORN, A.E.F., BANG, N.D., PEARCE, A.F., FLEMMING, B.W. & CARTER, R.A. 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. HEYMANS, J.J. & BAIRD, D. 1995. Energy flow in the Kromme estuarine ecosystem, St Francis Bay, South Africa. Estuarine, Coastal and Shelf Science 41: 39–59. HEEMSTRA, P. & HEEMSTRA, E. 2004. Coastal Fishes of Southern Africa. Grahamstown, NISC. 488 pp. JAMES, N.C. & HARRISON, T.D. 2008. A preliminary survey of the estuaries on the south coast of South Africa, Cape St Blaize, Mossel Bay – Robberg Peninsula, Plettenberg Bay, with particular reference to the fish fauna. Transactions of the Royal Society of South Africa 63: 111–127. JAMES, N.C. & HARRISON, T.D. 2009. A preliminary survey of the estuaries on the south coast of South Africa, Robberg Peninsula – Cape St Francis, with particular reference to the fish fauna. Transactions of the Royal Society of South Africa 64: 1–31. JERLING, H.L. & WOOLDRIDGE, T.H. 1991. Population dynamics and estimates of production for the calanoid copepod Pseudodiaptomus 84 Transactions of the Royal Society of South Africa hessei in a warm temperate estuary. Estuarine, Coastal and Shelf Science 33: 121–135. KOPKE, D. 1988. The climate of the Eastern Cape. In Bruton, M.N. & Gess, F.W. (Eds) Towards an Environmental Plan for the Eastern Cape. Grahamstown, Rhodes University. pp. 44–52. MARAIS, J.F.K. 1981. Seasonal abundance, distribution and catch per unit effort using gill-nets, of fishes in the Sundays estuary. South African Journal of Zoology 16: 144–150. MARAIS, J.F.K. 1983a. Seasonal abundance, distribution and catch per unit effort of fishes in the Krom estuary, South Africa. South African Journal of Zoology 18: 96–102. MARAIS, J.F.K. 1983b. Fish abundance and distribution in the Gamtoos estuary with notes on the effects of floods. South African Journal of Zoology 18: 103–109. MARAIS, J.F.K. 1984. Feeding ecology of major carnivorous fish from four eastern Cape estuaries. South African Journal of Zoology 19: 210–223. MARAIS, J.F.K. & BAIRD, D. 1980. Seasonal abundance, distribution, and catch per unit effort of fishes in the Swartkops estuary. South African Journal of Zoology 15: 66–71. MAREE, R.C., WHITFIELD, A.K. & BOOTH, A.J. 2000. Effect of water temperature on the biogeography of South African estuarine fish species associated with the subtropical/warm temperate subtraction zone. South African Journal of Science 96: 184–188 RATTE, T.W. 1989. Population structure, production, growth, reproduction and the ecology of Atherina breviceps Valenciennes, 1935 (Pisces: Atherinidae) and Gilchristella aestuaria (Gilchrist, 1914) (Pisces: Clupeidae), from two southern Cape coastal lakes. Unpublished PhD thesis. Port Elizabeth, University of Port Elizabeth. 332 pp. REDDERING, J.S.V. & ESTERHUYSEN, K. 1983. Sedimentation in the Kromme estuary. Port Elizabeth, University of Port Elizabeth ROSIE Report 6. 92 pp. REDDERING, J.S.V. & ESTERHUYSEN, K. 1984. Sedimentation in the Kabeljous estuary. Port Elizabeth, University of Port Elizabeth ROSIE Report 8. 42 pp. SCHLACHER, T.A. & WOOLDRIDGE, T.H. 1996. Axial zonation patterns of subtidal macrozoobenthos in the Gamtoos estuary, South Africa. Estuaries 19: 680–696. SCHARLER, U.M. & BAIRD, D. 2000. The effects of a single artificial freshwater release into the Kromme Estuary. 1: General description of the study area and physico-chemical responses. Water SA 26: 291–300. SCHARLER, U.M., BAIRD, D. & WINTER, P.E.D. 1997. Diversity and productivity of biotic communities in relation to freshwater inputs in three Eastern Cape estuaries. Pretoria. Water Research Commission Report 463/1/98. 197 pp. Vol. 65(1): 69–84, 2010 SKELTON, P.H. 1993. A Complete Guide to the Freshwater Fishes of Southern Africa. Halfway House, Southern Book Publishers. 388 pp. SMITH, M.M. & HEEMSTRA, P.C. 1991. Smiths’ Sea Fishes. Johannesburg, Southern Book Publishers. 1048 pp. STONE, A.W., WEAVER, A.B. & WEST, W.O. 1998. Climate and weather. In Lubke, R. & de Moor, I. (Eds) A Field Guide to the Eastern and Southern Cape Coasts. Cape Town, University of Cape Town Press. pp. 41–49 STRYDOM, N.A. & WHITFIELD, A.K. 2000. The effects of a single artificial freshwater release into the Kromme Estuary. 4: Larval fish response. Water SA 26: 319–328. TALBOT, M.M.J-F. 1982. Aspects of the ecology and biology of Gilchristella aestuarius (G & T) (Pisces: Clupeidae) in the Swartkops estuary, Port Elizabeth. Unpublished MSc. thesis. Port Elizabeth, University of Port Elizabeth. 128 pp. VORWERK, P.D., WHITFIELD, A.K., COWLEY, P.D. & PATERSON, A.W. 2003. The influence of selected environmental variables on fish assemblage structure in a range of southeast African estuaries. Environmental Biology of Fishes 66: 237–247. WALLACE, J.H. & VAN DER ELST, R.P. 1975. The estuarine fishes of the East Coast of South Africa, IV. Occurrence of juveniles in estuaries. V. Ecology, estuarine dependence and status. Durban. Investigational Report of the Oceanographic Research Institute 42. 1–63 pp. WALLACE, J.H., KOK, H., BECKLEY, L., BENNETT, B., BLABER, S.J.M. & WHITFIELD, A.K. 1984. South African estuaries and their importance to fishes. South African Journal of Science 80: 203–207. WHITFIELD, A.K. 1998. Biology and ecology of fishes in southern African estuaries. Icthyological Monograph of the J.L.B. Smith Institute of Ichthyology 2: 1–223. WHITFIELD, A.K. 2000. Available scientific information on individual South African estuarine systems. Unpublished Water Research Commission Report 577/3/00. 217 pp. WHITFIELD, A.K. & KOK, H.M. 1992. Recruitment of juvenile marine fishes into permanently open and seasonally open estuarine systems on the southern coast of South Africa. Ichthyological Bulletin of the J.L.B. Smith Institute of Ichthyology 57: 1–39. WHITFIELD, A.K. & HARRISON, T.D. 1996. Gilchristella aestuaria (Pisces: Clupeidae) biomass and consumption of zooplankton in the Sundays Estuary. South African Journal of Marine Science 17: 49–53. WOODWELL, G.M., RICH, P.H. & HALL, C.A.S. 1973. Carbon in estuaries. In Woodwell, G.M. & Peron, E.V. (Eds) Carbon and the Biosphere. Springfield VA, National Technical Information Service. pp. 221–240. WOOLDRIDGE, T.H. & MELVILLE-SMITH, R. 1979. Copepod succession in two South African estuaries. Journal of Plankton Research 1: 329–341.