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High Complexity Food Webs in Low- Diversity Eastern Pacific Reef-Coral Communities

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High Complexity Food Webs in Low- Diversity Eastern Pacific Reef-Coral Communities Ecosystems (2004) 7: 358-367 DOI: 10.1007/S10021-004-0184-X ECOSYSTEMS! ) 2004 Springer-Verlag High Complexity Food Webs in Low- diversity Eastern Pacific Reef-Coral Communities Peter W. Glynn Division of Marine Biology and Fisheries, Rosenstiel School of Marine and Atmospheric Science, 4600 Rickenbacker Causeway, Miami, Florida 33149, USA ABSTRACT Community-wide feeding interrelationships in a upwelling in the Pearl Islands results in high plank- low-diversity coral reef off the Pacific coast of Pan- ton productivity, which likely augments production amá (Uva Island reef) demonstrate complex path- in invertebrates, fishes, marine mammals, and sea- ways involving herbivore, strong corallivore, and birds, but these pathways still remain largely un- carnivore interactions. Four trophic levels with 31 quantified. The corallivore Acanthaster is absent interguild links are identified in a generalized food from upwelling centers in Panamá and from up- web, and documented feeding interrelationships welling and nonupwelling areas in the southern with 287+ species links are portrayed in a coral- and central Galápagos Islands, and the highly de- corallivore subweb. The importance of trophic structive, facultative corallivore Eucidaris galapagen- groups changes greatly with time, from unknown sis occurs only in the latter offshore islands and at causes over annual to decadal-scale periods, and Cocos Island. Relatively recent declines in the during very strong El Nino-Southern Oscillation abundances of manta rays, sharks, and spiny lob- events such that intermittent intense herbivory by sters are correlated with, but not necessarily caus- echinoids (Diadema) and corallivory by gastropod ally linked to, increasing fishing activities in the late moUusks, the crown-of-thorns sea star Acanthaster, 1970s to early 1980s. The extent to which the com- hermit crabs, and fishes result in high levels of coral plex yet highly unstable Uva Island food web is mortality and bioerosion of reef substratum. Intrar- representative of other eastern Pacific coral reef egional differences in species composition and ecosystems remains to be investigated. abundances affecting food-web interactions are briefly described for nonupwelling (Uva Island) and Key words: food web; coral reef; eastern Pacific; upwelling areas (Pearl Islands) in Panamá. Seasonal Panamá. INTRODUCTION ecosystems in the Indo-West Pacific and Greater Caribbean biogeographic regions support from two With only about 8-10 zooxanthellate (reef-build- to more than ten times the number of reef-building ing) coral species present on any given reef, and coral and fish species present on eastern Pacific three genera {Pocillopora, Porites, and Pavona) con- coral reefs (Paulay 1997). Numerous direct and in- tributing dominantly to reef accretion in the eastern direct feeding effects involving polychaetous Pacific, the array and complexity of known trophic worms, crustaceans, moUusks, echinoderms, and interactions are unexpectedly high in this low-di- fishes have been identified during the last 30 years versity coral reef region. High-diversity coral reef of coral community studies ranging from the Gulf of California to Ecuador (Barham and others 1973; Gilchrist 1985; Glynn 1973, 1974, 1976, 1977b, Received 10 May 2002; accepted 30 December 2002; published online 27 April 2004. 1982a, 1982b, 1983a, 1983b, 1984, 1985a; Glynn Corresponding author: e-mail: [email protected] and others 1979, 1982; Glynn and Colgan 1988; 358 Eastern Pacific Coral Reef Food Webs 359 70' conditions between the wet and dry seasons (Table Elevated sea temperatures associated with two exceptionally strong El Nino-Southern Oscillation '•V:^ Mexicci (ENSO) events in 1982-83 and 1997-98 resulted in Gulf of Califarnia extensive reef-coral mortalities over most of the equatorial eastern Pacific (Glynn 1990, 2002; Glynn and Colgan 1992, Glynn and others 2001). ENSO- induced coral mortality in the Galápagos was 97% Gulf of Çhiriqui ^ '^ and 26% following 1982-83 and 1997-98, respec- Uva Is. reef tively. The Uva reef suffered about 75% overall PACIFIC OCEAN Gulf of Panama coral mortality following the 1982-83 ENSO and about 13% from the 1997-98 ENSO. These distur- Galapagos Islands bances need to be considered because they signifi- cantly altered the relative abundances of corals, the numerous species that prey on them, and the im- portant links in the food-web design. It is also im- portant to note that the magnitude of fluxes varies 110° 100° 90° 80° 70° greatly over time, based on the presence of various species during a 30-year period. Several of these Figure 1 Eastern Pacific coral reef areas and location of species have demonstrated marked variations in Uva Island coral reef study site. abundance, intermittently contributing signifi- cantly to food-web interactions or, when rare, scarcely having any effect. Guzman 1988a, 1988b; Guzman and López 1991; Guzman and Robertson 1989; Reyes Bonilla and COMMUNITY-WIDE FOOD WEB Calderón Aguilera 1999; Wellington 1982a). How- ever, no comprehensive analysis of food-web rela- Four trophic levels with 31 interguild links (includ- tionships has yet been published for any coral reef ing at least five intraguild pathways) have been community in the eastern Pacific. identified in the Uva Island coral community (Fig- I construct the first ecosystem-level food web of a ure 2). All nonreferenced, generalized, and species- coral reef in Panamá that my colleagues and I have specific pathways presented here and below for continuously studied since 1970. This coral reef is macrobiota and fishes are based on personal con- located in the Gulf of Chiriqui (Figure 1), a nonup- sumer-prey observations and gut-content analyses performed on field populations. Detritus, constitut- welling environment supporting numerous small ing particulate organic matter (for example, decay- patch and fringing pocilloporid reefs. The reef is ing organisms and feces), receives contributions built predominantly by two frame-building coral from all reef biota and is consumed by decomposers species in the genus Pocillopora (P. damicornis and P. and by suspension and deposit feeders. Decompos- elegans), with secondary contributions from species ers such as microbes and species of meiofauna and in the genera Parités, Gardineroseris, Pavona, Psammo- microfauna are present in organic-rich sediment cora, and Millepora (a hydrocoral). Unlike coral reefs patches at the reef base, often encircling coral col- in other regions, where crustose coralline algae are onies and forming deposits in the internal structure often abundant and contribute importantly as bind- of reefs where they accumulate in the highly po- ing agents in framework construction, calcifying rous basal reef framework. The organic matter con- algae are generally unimportant to reef develop- tent of sediments is also enriched by particulate ment in the eastern Pacific. The Uva Island patch fallout from river runoff and plankton advected reef covers about 2.5 ha and consists of a reef flat, onto the reef. Studies of meiofauna species under- reef slope, and seaward reef base at a maximum taken on a Costa Rican reef found foraminifers, depth of approximately 6 m relative to mean low copepods, nematodes, and gastropods to predomi- water (MLW) (Glynn and Maté 1997; Eakin 2001). nate (Guzman and others 1987), and this is ex- Core drilling shows the reef framework to be 8-12 pected for the Uva Island reef. This rich (largely m thick, with a maximum age of about 5000 years microscopic) community demonstrates complex (Glynn and Macintyre 1977). Strong seasonal pat- feeding interactions involving meiofauna that prey terns at Uva Island greatly influence environmental on microbes, and microfauna that consume meio- 360 P. W. Glynn Table 1 Physical Environmental Conditions Influencing Offshore Coral Reefs in the Gulf of Chiriqui, Panamá Factor Wet Season Dry Season Cloud Cover" 23.2-41.7 6.8-9.2 median % Rainfall 423, maximum (Sep) 16, minimum (Feb) mm*', mean annual 2,598 Salinity, surface" 28.0 (27.0-32.0) 34.0 (28.0-36.0) Median (range) Tides^ low water exposures near low water exposures at midday midnight semidiurnal, unequal mean spring low salinity stress from rainfall intense solar exposure, heating, range = 3.29 m desiccation Light Penetration'' 10, minimum (Sep) 37, maximum (Jan) depth (m) of 10% surface radiation 0.5''-1.0'= (Jul-Sep) Sea Temperature (°C)''^ 30-31, maximum 25 minimum mean annual SST a~ 28°C (14 wk) (1 wk) Nutrient Concentration'' 0.3, minimum (Oct, Nov) 1.1, maximum (Mar) PO4-P (|xg-at 1"') at 40 m depth "Glynn 1977b ''Kwiecinski and Chial 1983 ^U.S. Department of Commerce, calculated from tidal predictions for Balboa, Panama ^Glynn unpublished observations 'Dana 1975 ^Renner 1963 ^Glynn and others 2001 fauna; hence, the redirected arrow in trophic cate- pendent on the availability of Zooplankton (> 95 gory 4 (Figure 2). |jLm net mesh size), whereas the branching coral Reef waters in the Gulf of Chiriqui are often filled Pocillopora damicornis grew independent of zoo- with phytoplankton and Zooplankton, depending plankton supply. upon nutricline shoaling and nutrients advected to Turf and crustose coralline algae are most abun- reefs via wet-season river discharge (L. D'Croz un- dant on reef fiats and in reef base habitats, whereas published data). The sporadic abundance of plank- green frondose algae occur most commonly in the ton populations and ill-defined current movements deeper reef base and talus slope zones. Reef slopes around the Uva reef make this potential food source in forereef and backreef zones are dominated by difficult to track and quantify. Phytoplankton and nearly continuous stands of live Pocillopora spp. (Ta- Zooplankton are consumed by suspension feeders, ble 2). The occurrence of macroalgae is intermittent such as sponges, sea anemones, polychaete worms, and may depend on nutrient concentrations arising lithophage bivalves, vermetid gastropods, barna- from pycnocline shoaling and rivers and runoff as cles, porcelanid crabs, and brittle stars, and by zoo- well as fiuctuations in herbivore populations. When plankton carnivores. These animals in turn contrib- abundant, macroalgae such as Caulerpa spp.
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