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Factors Affecting the Distribution, Abundance and Diversity of Rock-Pool Fishes on the Cape Peninsula, South Africa

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Factors Affecting the Distribution, Abundance and Diversity of Rock-Pool Fishes on the Cape Peninsula, South Africa Factors affecting the distribution, abundance and diversity of rock-pool fishes on the Cape Peninsula, South Africa B.A. Bennett and C.L. Griffiths Zoology Department, University of Cape Town, Rondebosch A quantitative description of the fish communities inhabiting rock Existing literature on the biology of intertidal fish has been pools on the Cape Peninsula, South Africa, and the relationships comprehensively reviewed by Gibson (1969, 1982) who points between the physical characteristics of the pools and the distribu­ out that relative to other elements of the intertidal community, tion, abundance and diversity of the resident fish fauna are given. fish have received remarkably little attention. Most studies on A total of 1 541 fish, representing 21 species, were collected from 84 rock pools using the ichthyocide Rotenone. The mean fish have, moreover, dealt with the northern hemisphere while 2 2 density was 7,42 fish m- (49,60 9 m- ) of pool area or 0,58 fish African, Australian and South American shores remain poorly m -2 (3,67 9 m -2) of the whole intertidal zone in which the pools known. were situated. Clinus superciliosus, C. cottoides and Chorisochis­ General information on the species composition, reiatiVt: mus dentex were the most abundant species, together comprising abundance and intertidal zonation of fishes inhabiting rock 75% of the biomass and 60% of the number of fish caught. The number of species, number of individuals and biomass of pools in the south-western Cape Province, South Africa, is fish in the pools all correlated significantly (P < 0,01) with pool given by Jackson (1950) and Penrith (1965). Neither report size and the amount of rock cover available. In addition, the was published, nor provided measures of absolute abundance. number of species decreased up the shore. These three environ­ Factors regulating density and species composition of rock­ mental factors in combination accounted for between 58% and . pool fish communities in South Africa and elsewhere remain ) 88% of the observed variance in diversity, number and biomass of 0 virtually unknown (Gibson 1982). Seasonal changes in diversity 1 fish in the pools. Rock cover was the single most important 0 variable, explaining up to 76% of the variance. Possible reasons occur and can be correlated with seasonal fluctuations in 2 for the importance of rock cover and its relationship with pool temperature, wave action and upwelling intensity (Green 1971; d e t size, are discussed. Grossman 1982; Thomson & Lehner 1976)_ South-western a S. Afr. J. Zool. 1984, 19: 97 -104 d Cape rock-pool fish populations remain stable in the long and ( r short term (Jackson 1950; Penrith 1965, 1970) perhaps because e 'n Kwantitatiewe beskrywing van die visgemeenskappe wat in die h there is little seasonal variation in water temperature (Griffiths s rotspoele van die Kaapse Skiereiland voorkom, word gegee en die i l 1977; Penrith 1970). b verhoudings tussen die fisiese eienskappe van die poele en die u verspreiding, getalsterktes en diversiteit van die visfauna is In this study, we provide the first quantitative estimates of P ondersoek. e the abundance and composition of South African intertidal h 'n Totaal van 1 541 vi sse van 21 spesies is uit 84 rotspoele met t fish communities and elucidate the physical attributes of rock y behulp van die visgif Rotenone gekollekteer. Die gemiddelde digt­ pools that influence the number and species of fish. The feeding b 2 2 heid was 7,42 visse m- (49,60 9 m- ) van die poel-area, of 0,58 d 2 habits of the fish collected during this study are reported e visse m- (3,67 9 m-1 van die hele getysone waarin die poele t elsewhere (Bennett, Griffiths & Penrith 1983). n voorkom. Clinus superci/iosus, C. cotto/des en Chorisochismus a r dentex was die volopste spesies en het saam 75% van die g biomassa en 60% van die getal visse wat gevang is, gevorm. Methods e c Die aantal spesies, aantal individue en biomassa van die visse Samples were collected from six intertidal localities around the n e in die rotspoele het betekenisvol gekorreleer (P < 0,01) met poel­ c Cape Peninsula, South Africa; three in False Bay and three i l groolle en die hoeveelheid rotsbedekking beskikbaar. Daarby het r die aantal visse teen die strand op verminder. Hierdie drie omge­ along the West Coast (Figure O. Each site was visited at low e d wingsfaktore saam was verantwoordelik vir tussen 58% en 88% water of spring-tide (L WS) between July and September 1982 n van die variasie wat waargeneem is in die diversiteit, getalle en and pools representative of the full height and size range were u y biomassa van die vis in die poe Ie. Die belangrikste enkele veran­ selected. The height (HGH1) and distance (DIS1) of each pool a derlike was rotsbedekking wat tot 76% van die variasie kon ver­ w from the L WS mark were recorded and its area (AREA), e duidelik. Moontlike redes vir die belangrikheid van rotsbedekking t maximum depth (MAXD) and mean depth were measured. a en die verhouding daarvan met poelgroolle word bespreek. G Pool volume (VOL) was calculated by multiplying mean depth S.-Afr. Tydskr. Dier/(. 1984, 19: 97 -104 t e by area. Subjective ratings of rock (ROCK) and algal (WEED) n i cover available as shelter to the fish were made on a scale of b a zero to ten. Total pool area (including pools that were not S y B.A. Bennett· and C.L. Griffiths sampled) in 20 m wide transects was measured and expressed b Zoology Department, University of Cape Town, Rondebosch, as a percentage of the total area included in each transect. d e 7700 Republic of South Africa c °To whom correspondence should be addressed After Rotenone (dissolved in acetone) had been added to u d the pools, fish were collected with hand-nets and immediately o r Received 18 July 1983; accepted 6 October 1983 transferred to 10070 formalin. Each fish was subsequently iden- p e R 98 S.-Afr. Tydskr. Dierk. 1984, 19(2) log (independent variable transformed) and log-log models were used for two reasons: firstly, the data were in some cases not normally distributed and secondly, to detect possible non­ linear relationships between the variables. + Unless otherwise stated, where pairs of figures are given in GRANGER BAY. the text, the order is always False Bay: West Coast. THREE ANCHOR BAY_ BEA POINT- Results and Discussion The fish communities A total of 1 541 fish representing 21 species were collected from 34°8 84 rock pools (Table 1). Sampling effort was greater on the West Coast where 1 028 fish with a biomass of 6286 g were collected from 50 pools. In False Bay, 34 pools yielded 513 fish with a mass of 4 021 g. Population densities on the two sides of the Peninsula were very similar, both numerically (6,82 and 7,75 fish m-2 pool area) and in terms of biomass (53,45 and 47,41 g m-2 pool area). When calculated in terms of the whole shore area in which the pools were situated, densities 2 FALSE BAY were again similar: 0,55 and 0,60 fish m- ; 4,30 and 3,20 g 2 ATLANTIC m- • OCEAN It is very difficult to compare these figures with any of the previously published information on the densities of rocky shore fish populations elsewhere. Gibson (1982) provides a table of such figures and discusses the problems facing dif­ ferent authors in their choice of units for expressing density. In view of the lack of available information concerning fac­ tors as the number of pools per unit shore area, pool size CAPE POINT 1 250000 distribution, the proportion of fish inhabiting areas other than ~ ..•• ~ .... 1~ Km 1S030'W rock pools and the proportions of cryptic as opposed to free­ swimming species, detailed comparisons are not possible. It . Figure 1 Cape Peninsula, South Africa, showing the six sites from which suffices to say that Gibson's (1982) observation that 'densities ) 0 rock-pool fish were sampled. rarely exceed a few individuals per square metre' also holds 1 0 true for the south-western Cape. 2 d Species distribution and abundance varied considerably be­ e t tified to species according to Pemith (1969) for clinids and tween the two sides of the Peninsula. Eight species were com­ a d Smith (1965) for other species, measured to the closest milli­ mon to both sides with eight restricted to False Bay and five ( r metre (total length) and weighed to within 0,01 g (wet mass). to the West Coast (Table 1). The greater number of species e h in the False Bay sample (despite the smaller sampling effort) s i l Data analysis reflects the general trend in diversity of intertidal species be­ b u Simple linear regression analysis was performed on all possible tween the two sides of the Peninsula (Stephenson 1939, 1944, P e two-way combinations of dependent variables (number of 1948) and has been demonstrated in the Clinidae by Pemith h t species, number of fish and biomass) and independent variables (1970). y Abundance values for False Bay, for the West Coast and b (HGHT, DIST, AREA, VOL, MAXD, ROCK and WEED). d The respective correlation coefficients were tested for for the combined data are shown in Table 1. Clinus cottoides e t significance (Ho : r = 0; HI : r '#- 0) and bivariate scatter plots was numerically the most important species on the False Bay n a coast, followed by Caffrogobius coffer, Clinus superciliosus r were examined to detect strongly deviating data.
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