Spatial Patterns of Bicolor Damselfish Populations in Jamaica and St
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SPATIAL PATTERNS OF BICOLOR DAMSELFISH POPULATIONS IN JAMAICA AND ST. CROIX ARE DETERMINED BY SIMILAR POST-SETTLEMENT PROCESSES R. S. Nemeth Department of Zoology, University of New Hampshire, Durham, NH 03824 [Converted to electronic format by Damon J. Gomez (NOAA/RSMAS) in 2003. Copy available at the NOAA Miami Regional Library. Minor editorial changes were made.] Proc 8th Int Coral Reef Sym 1 :1017-1022 . 1997 SPATIAL PATTERNS OF BICOLOR DAMSELFISH POPULATIONS IN JAMAICA AND ST . CROIX ARE DETERMINED BY SIMILAR POST-SETTLEMENT PROCESSES R .S . Nemeth Department of Zoology, University of New Hampshire, Durham, NH 03824 USA ABSTRACT Itzkowitz 1977 ; Waldner and Robertson 1980) . Adults occupy Porites porites and Montastrea annularis Larval supply, larval behavior and post-settlement colonies, sponges and large rubble but are largely processes all affect the abundance and distribution of restricted to fore reef habitats . This unique size- marine fish populations . Post-settlement processes were class distribution was examined with recruitment surveys found to largely determine distribution patterns of and with experiments measuring the variation in juvenile Although S . feeding behavior, competitive interactions, growth and bicolor damselfish (Stegastes partitus) . partitus larvae settle at similar densities to both back survival among different habitats and substrates . The reef (BR) and fore reef (FR) habitats, adults are more demographic responses to these conditions were measured abundant on the FR . I tested the hypothesis that on two islands in the Caribbean to examine the similar post-settlement processes were acting among generality of the results at a larger spatial scale . distant sites to produce this general pattern of distribution and population structure . Juvenile mortality was significantly greater on Montastrea MATERIALS AND METHODS Porites rubble . This effect was annularis coral than on similar among FR and BR habitats and between St . Croix This study was conducted in Teague Bay, St . Croix, and Jamaica . The effects of substrate type on growth United States Virgin Islands (17 ° 45' N, 64 ° 42' W) and was minor with some suggestion that juveniles grow Discovery Bay, Jamaica (18 ° 27' N, 77°24' W) . Since the faster on Porites rubble . Increased competition for population structure of Stegastes partitus had not been space and decreased planktonic food reduced juvenile previously documented in St . Croix, juvenile and adult . S . densities were visually sampled in back reef (BR) and growth rates in BR habitats on both islands partitus living in the BR take twice as long to reach fore reef (FR) habitats during July from 1991 to 1994 . adult size, making them more vulnerable to predation Juvenile and adult fish were counted along 30 x 1 m during this prolonged juvenile period . The combined strip transects (n=16) by divers who held a 1 m wide T- effects of these factors are sufficient to reduce adult shaped bar for reference . Census data on juvenile and abundance on the BR thereby producing the spatial adult abundance in BR and FR habitats were averaged for patterns of distribution observed among reef zones in each year (n=4) and analyzed with ANOVA . this and other studies in the Caribbean . The effect of different substrate types within each habitat on juvenile growth and survival was examined in INTRODUCTION Teague Bay in August 1992 using an orthogonal experimental design . I selected four isolated M . The dispersive larval stage of marine organisms annularis coral heads (1 to 1 .25 m 2 ) and constructed functions to colonize patchily distributed habitats or four P . porites rubble piles (1 m 2 ) in sand flats in unpredictable environments while spreading the risk of . adjacent FR (10-15 m depth) and BR (1-5 m) habitats local extinction across a wide geographical range . partitus were collected and marked with Juvenile S (Barlow 1981) . This trait has focused attention on the tattoo ink and treated in a 0 .25 g/l bath of importance of variation in the arrival of colonizing tetracycline hydrochloride for 12 hr to mark their individuals in determining the distribution and . After the 16 experimental otoliths (Hettler 1984) abundance of reef fishes (Doherty and Williams 1988 ; units were denuded of all fishes, the tagged juveniles Doherty and Fowler 1994) . Alternative explanations have were stocked at 4 fish/m 2 . Fish length averaged 14 .1 mm considered the effects of processes occurring after settlement in structuring populations (Smith and Tyler SL (-1 .73 SD) . During the experiment recruitment of . 1978 ; Gladfelter et al . 1980) . other species was allowed to occur . The area around the 1972 ; Talbot et al Recent evidence indicates that the interactions between study site was also censused twice for possible S . fishes and their relationship within the physical and partitus emigrants . In only two instances, once on the BR and once on the FR, was an experimental fish found on biological environment may prove to be important in predicting changes in population structure (Eckert 1985 ; an adjacent coral head . This occurred within the first Jones 1988, 1991 ; Forrester 1990, 1995 ; Hixon 1991 ; two days after stocking so each fish was captured and . Robertson 1996) . returned to the nearest experimental unit, a coral head Potential predators occupying or visiting experimental When a larva makes the transition to the reef units were counted and these data were used to estimate environment, its fitness will be increased if it selects predator densities in FR and BR study sites . a place to live where predation is minimized and growth is maximized (Gilliam and Fraser 1987) . While patterns Feeding behavior (bites/min in plankton and benthos) and aggressive interactions (chases/min) were recorded for in juvenile distribution, growth and mortality have been ; each fish during three 10 min observation periods . detected in a number of reef fishes (Jones 1986,1988 Wellington 1992), the habitat characteristics Aggressive encounters were grouped as intraspecific aggression (adult + juvenile S . partitus), interspecific responsible for these patterns and their relative importance on subsequent population structure remain aggression (all other Stegastes spp .) or aggression with ; Richards and Lindeman 1987) . other non-pomacentrid spp . A total of 57 hours of unknown (Hadfield 1986 The extent to which habitat characteristics influence observations were recorded from 22 FR fish and 20 BR fish between 15 and 26 August from 08 :30 to 18 :30 . At population structure requires information on fish- habitat associations . For example, in a group of six the end of the two week study, surviving fish were collected and their otoliths extracted . Lapillar ecologically similar Atlantic pomacentrids of the genus Stegastes, the juveniles and adults display habitat otoliths were viewed under UV light and the distance from the tetracycline mark to the edge of the otolith distribution patterns which are fairly consistent ; Clarke 1977 ; was used as a measure of fish growth . Since lapillus throughout the Caribbean (Emery 1973 Itzkowitz 1977 ; Waldner and Robertson 1980) . The radius (LR) was highly correlated with standard length mechanisms generating these patterns at one location may of fish (r 2 =0 .90, P<0 .001), it was used to calculate include habitat selection at settlement, differential individual growth rate in standard length (SL) with the mortality after settlement, or unequal competitive regression equation : SL = -1 .776 + 56 .1(LR) . The abilities among species (Wellington 1992, Robertson formation of daily growth increments on otoliths has 1996) . been experimentally verified for S . partitus by (Robertson et al . 1988) . Growth and behavioral data In this study I examine the importance of these were averaged to yield a single datum point per mechanisms in affecting the distribution and population experimental unit and analyzed using two-way analysis of structure of Stegastes partitus Posy (Pomacentridae) . covariance with zone (FR v s . B R ) and substrate type Juveniles primarily occupy rubble substrates on back (live coral vs . coral rubble) as fixed factors and reef and fore reef habitats (Emery 1973 ; Clarke 1977 ; standard length as a covariate . Prior to analysis data 1018 Nemeth were tested for homogeneity of variances using more on planktonic organisms than fish living on rubble Bartlett's chi-square (Sokal and Rohlf 1981) . When in both BR (34% vs . 16%) and FR (59% vs . 17%) habitats . necessary data were transformed ln(X + 1) to meet Intraspecific aggression was greater for juveniles assumptions of homoscedasticity . living on the FR due to the presence of several adult conspecifics which moved onto experimental units . In St . Croix the relative supply of planktonic organisms Conversely, interspecific aggression was higher on the available to juvenile S . partitus in BR and FR habitats BR due to recruitment of S . leucostictus but also S . was measured using tube traps (Yund et al . 1991) . The planifrons . Aggression with non-pomacentrid species was clear, cylindrical acrylic tubes (60 cm height x 5 .1 cm similar among substrates and habitats (Table 2) . inside diameter) were filled with a 10% solution of buffered (borax) formaldehyde and sea water with a few In Jamaica, patterns of growth of juvenile S . partitus drops of food color added to monitor formalin level . matched those found in St . Croix . Fish living on the FR Twelve tube traps were deployed in pairs at each site . had faster growth rates than fish on the BR with no At the end of the study, tube traps were retrieved, difference among substrate types (Fig . 2) . Moreover, filtered through a 75 gm mesh sieve and the preserved juvenile S . partitus showed depressed growth rates planktonic organisms from each tube were counted and living in the presence of the territorial competitor, S . classified to ordinal level . leucostictus (0 .105 --0+ .008 vs . 0 .143--0+ .011 mm/d SL, .59, P<0 .002( . Based on the results of this study, a similar experiment F1,21=12 was conducted in Discovery Bay, Jamaica in February Fish survivorship was significantly higher on Porites 1994 .