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The Influence of Post-Settlement Processes on the Structure of Reef-Fish Assemblages

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The Influence of Post-Settlement Processes on the Structure of Reef-Fish Assemblages The influence of post-settlement processes on the structure of reef-fish assemblages Anne-Marie Eklund, Alina M. S2mant, and James A. Pohnsack Running head: Post-§ettlement processes affecting reef fish assemblages The influence of post-settlement processes on the structure of reef-fish assemblages ABSTRACT We used model reefs with varying amounts of habitat complexity and epibenthic growth to test the effects of reef shelter and food resources on fish density, species richness, biomass, size composition and species relative abundances. We visually censused the reef fish assemblages on each model reef, monthly from September 1991 through December 1993, and estimated fish biomass through length/weight regressions from fish collections. Reefs with different amounts of reef-based food resources did not support significantly different fish assemblages; however, reef shelter had a significant effect on the fish assemblage structure. High-shelter reefs sustained greater fish densities, but low-shelter 49 reefs supported greater fish biomass. The size and species composition varied between reef types with a greater abundance of juvenile fishes on the reefs with more shelter from predators. Although juvenile fishes less than 2 cm TL settled indiscriminately on all of the model reefs, they did not persist on reefs lacking adequate shelter. Fishes from 2-1 0 cm TL were. significantly more abundant on high-shelter reefs. The tightly coupled post-settlement processes of predation and competition for limited shelter from predators greatly affected the fish assemblage structure. Post-settlement predation and resource limitation, therefore, may be more important than larval supply in regulating Caribbean/Atlantic reef fish assemblages. I INTRODUCTION Coral reefs are extremely complex habitats which support a high density and diversity of fishes. The mechanism(s) responsible for maintaining dense and speciose assemblages have been debated by many reef fish ecologists (e.g. see Doherty and Williams, 1988; Hixon, 1991; Jones, 1991; Sale, 1991). Determining the limiting factors of reef fish production is of interest to fishery managers and ecologists alike. For the past 20 years, there has been a greater emphasis in the literature on the stochastic and highly variable planktonic environment and its effects on larval fish survival and recruitment. Several studies have concluded that reef fish populations appear to be recruitment limited, which means that they are limited by processes that occur during the larval planktonic stages (Talbot et al., 1978; Victor, 1983,1986; Doherty and Williams, 1988; Doherty and Fowler, 1994a,b). There is some evidence (Doherty and Williams 1988; Doherty and Fowler, 1994a,b) that larval mortality rates are sufficiently high to reduce population numbers to the point where reef resources are not limiting. If population numbers are very low, then the amount of shelter or food on coral reefs would not be limited for reef fishes, and therefore, they would not be important variables in controlling fish populations and the resulting reef fish assemblage structure. Although some species of pomacentrids on the Great Barrier Reef may be recruitment limited (Doherty and Fowler, 1994a,b), there is evidence that some families of fishes in the Atlantic (e.g. Haemulidae) may be subject to high rates of post-settlcment predation (Shulman and Ogden, 1987; Hixon and Beets, 1993). The amount of shelter available to those reef fishes may have a direct effect on their survival. Post-settlement predation may 2 be high enough io have an effect on overall population sizes and species relative abundances. It is possible to assess the importance of reef resources in controlling fish assemblage structure and production by manipulating the amount of reef shelter or food available. If a . reef fish population is limited by habitat, then a complex reef, offering more shelter, could support more fish, particularly small prey fish. Shulman (1984) found a significant relationship between shelter availability and the abundance of small prey fishes in the Virgin Islands. The applicability of her experiments to large reef systems has been questioned, however, due to the very small scale of the model reefs that she used (conch shells) (Doherty and Williams, 1988). Hixon and Beets (1989, 1993) also studied the relationship between reef shelter and densities of different groups of fishes on artificial reefs in the Virgin Islands, but their results were inconsistent, possibly due to seasonal recruitment pulses. Neither Shulman (1984) nor Hixon and Beets (1989,1993) examined changes in fish biomass as an indicator of fish production. By following patterns of fish density, species richness and biomass on larger experimental reefs monthly for over two years, we have been able to substantiate and expand upon the previous studies' conclusions. To test whether increased shelter availability does increase reef fish populations, we compared model reefs having identical external structures but very different internal structures - low shelter (hollow) vs. high shelter (filled) reefs. Similarly, to test for the effect of reef food resources on reef fish populations, we compared reefs with identical structures but with different amounts of epibenthic growth. We measured changes in fish density, biomass, species richness and size and species composition on the different reef types. Our hypothesis was that fish density, biomass and species richness on high shelter or high food 3 reefs would be greater than on low shelter or low food reefs and that the frequency distribution of fish by species and by size class would differ among reefs with different shelter or food resources. By looking at fishes of different species and size classes, it was possible to determine if some target species are sensitive to changes in habitat quality or if the juvenile stages of some species are limited by habitat. This study improves upon previous ones, such as Shulman (1984) and 1-fixon and Beets (1989, 1993), by examining the ontogeny of shelter dependence, by conducting the study over a larger area and longer time period, and by further defining the relationships of different size classes and species to shelter resources. METHODS Study Site and Treatments Twelve prefabricated concrete model reefs were used to provide four treatments of habitat complexity and food resources (three replicates per treatment). The reefs were deployed in 9 in of water ca. 500 in off the coast of Palm Beach, Florida in August 1991 (Figure I The pyramid shaped modules were 2.4 rn^ at the base, I in' at the truncated top (which was open) and 1.8 in high (Figure 1). Along each of the four sides were 20 cm^ holes spaced 15 cm apart. The replicate reefs were placed 15 m apart along east-west transects, and each treatment was at least 400 in apart along a north-south axis parallel to the shoreline. The nearest natural reef areas were at least 250 in from the artificial structures. To test for the effects of vay*g reef food resources on the fish assemblages, we attempted to manipulate epibenthic growth. One set of reefs (FF) was fertilized with 4 Osmocote(& fertilizer to increase growth of attached epiflora. A second set (PF) was painted with anti-fouling paint to reduce the amount of epibenthic growth on the exterior surfaces. A third set (CF) was not manipulated. All three of these reef sets were filled with large pieces of broken cinder blocks. The blocks were placed inside the pyramids haphazardly, creating interstices of many sizes for small, cryptic fishes to inhabit. The reefs of the remaining set were not filled with cinder blocks and remained hollow structures (H). The hollow and filled reefs were compared to test for effects of reef shelter on the fish assemblages. Fish Censuses Fish density, size frequency, and species composition were determined through monthly visual censuses at each reef. An open sandy site between the fertilized and the control reef treatments was also censused in order to compare the reef fish assemblages with those on non-reef areas. The visual survey method was adapted from Bohnsack (1979), in which all fish on each reef are counted and measured by scuba divers. PVC rods with 30 cm rulers attached enabled divers to measure individual fish. Divers counted and measured all of the highly mobile species first, followed by closely examining the reef holes and crevices for the cryptic species. These surveys were supplemented by a night dive, so that the nocturnally active species could be observed. Because these model reefs are small isolated structures, it was possible to count and measure all of the fish seen, precluding the use of other visual techniques which "subsample" fishes on larger natural reefs. Fish Biomas In September 1993, one reef from each treatment was poisoned with rotenone, and 5 all fish were collected and measured for the development of length/weight relationships for each species. Prior to the rotenone sampling, each reef was censused visually in order to determine the biases or limitations of the visual censuses. Divers then encircled each reef with a 4.6 in radius cast net, by beginning upstream and slowly swimming downstream towards the reef with the outstretched net. In this way, those schooling species,that fed upstream from the reef (e.g. haemulids) were corralled as they sought the shelter of the reef. A 0.65% solution of pure rotenone was quickly sprayed into reef crevices, and divers collected fish as they expired. One diver remained above the reef with a net to intercept those very small fish that escaped through the cast net's mesh. Lengthtweight regressions for each species are listed in Table 1. Natural Reefs Assessments of representative fish assemblages for the area were done by censusing fish density and species composition on patch reefs within 500 rn of the model reef sites. The stationary census method used for the natural reefs was taken from Bohnsack and Bannerot (1986), where all of the fish observed in 5 minutes were identified, counted and measured.
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