Exploring the behavioral and reproductive ecology of fishes on coral reefs The 'lab rat' of coral reef fish research:
Thalassoma bifasciatum
Much of my research for the last decade has focused on herbivorous coral-reef fishes, particularly the Caribbean striped parrotfish, Scarus iserti. These relatively small, sex-changing, herbivores display a remarkably complex social organization (including group territories with dominance hierarchies, roving "satellite" individuals, and non-territorial schools that invade territories). Their range of mating tactics and alternative life-history strategies, coupled with their quantifiable food resources make them, in many ways, an ideal study organism. Additionally, the ability of these fish to respond quickly to changing environmental and demographic conditions makes them especially well-suited for both comparative and experimental field studies. By combining data on resource availability with experimental and comparative data on their behavior and demography my studies explore how physical and social environmental effects determine where animals are found and how they interact.
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My research to date includes studies of the economics of group-territorial behavior by non-kin (Clifton 1989, 1990), the effects of seasonal changes in resource levels upon social organization and the timing of reproduction (Clifton 1995), the determinants of group size inside and out of territories (Clifton, submitted), and tests of "ideal despotic" models of dispersion under variable regimes of resource renewal (in prep). Social behaviors such as dominance (Clifton 1991) and territoriality play a particularly important role in determining the dispersion and life history of these fish because they modify both individual food intakes and underlying patterns of food availability. |
The Beaugregory Damselfish Stegastes leucostictus (with white paint tag on its back) More recently, my studies of reef-fish reproductive phenology, in collaboration with Dr. Ross Robertson of the Smithsonian Tropical Research Institute, have investigated how food limitation constrains the life history, parental care, and reproductive ecology of the Caribbean damselfish (Stegastes leucostictus), a territorial herbivore with male parental care of eggs. In a series of field experiments, we supplemented the diet of wild adult fish over extended periods of time to judge how food availability influenced adult life history. Simultaneously, we monitored unmanipulated populations on a variety of spatial scales to establish the extent and relative importance of these effects. After almost two years of field work, we have clearly documented how food availability drives seasonal, but not lunar, patterns of reproduction, growth, and survival for this tropical species. This link between aspects of the adult environment and their expression of various life history traits challenges current thinking about the seasonality of reproduction on coral reefs and emphasizes the need to include adult biology within current models of reef fish population ecology. Plans to extend this work to other fishes with different life history and ecology (e.g. planktivores and detritivores without parental care) are currently underway.
Other Studies of Reef Fish Phylogenetic, ecological, and biogeographic relationships among Labroid coral reef fishes
Although relatively recently derived, parrotfishes and wrasses have rapidly evolved into one of the most diverse and influential guilds of fishes on modern coral reefs. Similarities in morphology, fin structure, and scale pattern have traditionally obscured the phylogenetic relationships within these important groups. In collaboration with a molecular population biologist (G. Bernardi at UCSC) we have recently established a molecular phylogeny for both the Atlantic parrotfish genus Sparisoma (Bernardi et al., submitted) and the pan-tropically distributed wrasse genus, Thalassoma (Bernardi et al., in prep). Levels of genetic similarity within Sparisoma provide intriguing insights into their molecular, ecological, and biogeographic divergence. Similarly, our on going analysis of Thalassoma phylogeny is revealing unexpected relationships that both challenge and support current perceptions of speciation within the group. Our success thus far prompts further study, and we are currently working toward a complete phylogeny of the Scaridae that examines patterns of evolution, ecology, and biogeography.
Diel feeding cycles by herbivorous fishes. Many herbivorous fishes show marked daily cycles in their feeding rate. Many are also territorial. As a result, several workers have suggested that feeding periodicity is a response to predictable changes in food quality (i.e. nutritive value) and the ability to control access to that food. A simple model I have developed, however, predicts that food quantity (i.e. standing crop) and competition for resources, may also generate feeding cycles. Ross Robertson and I are discriminating among these possibilities using a comparative and experimental approach. To date, we have characterized the diel feeding behavior of 13 species of herbivorous fish (damselfish, surgeonfish, and parrotfish). Together, these fish show a range of feeding patterns, express different levels of territorial behavior, and eat different types of algae and sea grass. A preliminary analysis indicates that various factors drive feeding cycles in different species, depending on resource type, demography, and digestive physiology. Experimental manipulations of food quality, food quantity, and population density will test the models further.
Relevant publications Clifton, K.E. & Robertson, D.R. (in prep). Diet and reproduction of the Caribbean Yellow jack, Carangoides bartholomaei in the near-shore waters of San Blas, Panama.
Robertson, D.R. & Clifton, K.E. (in prep). Patterns of mortality in the herbivorous Caribbean damselfish, Stegastes leucostictus.
Clifton, K.E. & Rogers, L. 2008. Sex-specific mortality explains non-sex-change by large female Sparisoma radians, a protogynous Caribbean parrotfish. Animal Behavior, 75:e1-10.
Bernardi, G., Clifton, K.E., Robertson, D.R. & Azzurro. E., 2000 Molecular sytematics, zoogeopraphy, and evolutionary ecology of the Atlantic parrotfish genus Sparisoma. Mol. Phyl Evol 15:292-300.
Clifton, K.E. & Clifton, L.M. 1998 A survey of fishes from various habitats within the Cayos Cochinos Marine Preserve, Honduras. Rev Biol. Trop. 46: 109-124.
Clifton, K.E., Kim, K. & Wulff, J.L. 1997 A field guide to the reefs of Caribbean Panama emphasizing western San Blas, Panama. Proc. 8th Int. Coral Reef Symp. 1: 167-184.
Clifton, K.E. 1996. Field methods for the behavioral study of foraging ecology and life history of herbivorous coral reef fishes. Proc. Am. Acad. Und. Sci. 16: 75-82.
Clifton, K.E. 1995. Asynchronous food availability on neighboring Caribbean coral reefs determines seasonal patterns of growth and reproduction for an herbivorous parrotfish. Mar. Ecol. Prog. Ser. 116:39-46.
Clifton, K.E. & Robertson, D.R. 1993. Risks of alternative mating tactics. Nature 366:520.
Clifton, K.E. 1991. Subordinate group members act as food finders within striped parrotfish territories. J. Ex. Mar. Bio. Ecol. 145:141-148.
Clifton, K.E. 1990. The costs and benefits of territory sharing for the Caribbean coral reef fish, Scarus iserti. Behav. Ecol. Soc. 26:139-147.
Clifton, K.E. 1989. Territory sharing by the Caribbean striped parrotfish, Scarus iserti, patterns of resource distribution, group size, and behaviour. Anim. Behav. 37:90-103.
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