MARINE ECOLOGY PROGRESS SERIES Vol. 160: 291-293, 1997 Published December 15 Mar Ecol Prog Ser NOTE Fish predation on the scleractinian coral Madracis mirabilis controls its depth distribution in the Florida Keys, USA A. G. Grottoli-Everett*, G. M. Wellington Department of Biology and Biochemistry, University of Houston, Houston, Texas, 77204-5513, USA ABSTRACT We evaluated the role of f~shpredation as the mann 1993). It forms densely packed clumps of small mechanism controlling the local distribution of the scleractin- pencil-sized branches with blunt tips, small polyps ]an coral Madracis mirabills on Conch Reef in the Florlda which are light brown to yellow in appearance. M. Keys, USA At many Car~bbeansltes, M m~rabilisis found at depths ranging from 1 to 63 m However, at Conch Reef th~s mirabilis is found nearshore at a depth of 3 m at the coral IS rarely found shallower than 20 m M ~nlrabilisfrag- Channel Two pass (24"50' 05" N, 81'41' 05" W) which ments transplanted from 26 to 13 m depth suffered sign~fi- links Florida Bay and the Florida reef track between cantly hlgher levels of f~shpredation than fragments trans- Lower Matacumbe Key and Long Key (pers. obs.). At planted from 26 to 20 m These results Indicate that fish predat~onpotentially 11mits the ve~tlcal distribution of M Conch Reef, Florida (24"57' 05" N, 80" 28' 00" W), it is mlrabllis on Canbbean reefs rarely found shallower than 20 m. In the course of experimentally assessing the source of carbon in the KEY WORDS: Fish predat~on- Col-al distnbution Madracis skeletal matrix of M. mirabihs, we observed evidence mlra biljs of heavy fish grazing on colonies transplanted from 26 to 13 m depth. In this experiment, 8 distinct Madracis mirabilis coral colonies were collected at 26 m and each was Competition for space, habitat type, food availability, divided into 8 fragments. Each individual fragment environmental conditions and natural disturbances was cemented to an individual plexiglass disk. Four have often been emphasized as important factors con- fragments from each coral head were transplanted to trolling coral distribution (Loya 1972, Porter 1972, 13 m and 4 to 20 m (4 fragments colony-' X 2 depths X 1974, 1976, Fishelson 1973, Lang 1973, Glynn 1976, 8 colonies = 64 total sample size). All fragments were 1988, Sammarco 1980, 1982, Wellington 1982a, Huston manipulated in an identical fashion at both depths. 1985, Glynn & Colgan 1988). Increasing evidence from Coral fragments were collected approximately 6 wk field studies shows that predation by invertebrates later and treated in 5% sodium hypochlorite solution (Robertson 1970, Glynn et al. 1972, 1979, Glynn 1976, for 24 h. Because the number of branches per fragment McClanahan & Shafir 1990, McClanahan et al. 1996) was unequal, predation is reported here as a percent- and corallivorous reef fish (Glynn et al. 1972, Kaufman age, as follows: 1977, Brock 1979, Neudecker 1979, Wellington 1982b, total number of fish bites Harmelin-Vivien & Bouchon-Navaro 1983, Littler et al. X 100% (total number of branches/fragment) 1989) can play a direct role in shaping coral distribu- tion and community structure (reviewed by Glynn In cases where the total number of bites exceeded the 1990, Hixon 1997).Here, we present evidence that fish total number of branches (due to more than 1 bite/ predation limits vertical distribution of the pocilloporid branch-') the percent predation value exceeded 100. coral Madracis mirabilis on the Florida, USA reef tract. Overall mean percent predation was found to be sig- Madracis mirabilis is commonly found throughout nificantly greater at 13 m than at 20 m (Table 1).With the Caribbean, South Florida and the Bahamas (Hu- the exception of colony 2, the mean percent predation was consistently higher in shallow water. In some shal- low water fragments, all branch tips were bitten off, yet none of the fragments in deeper water were in such 0 lnter-Research 1997 Resale of full artjcle not pernutted Mar Ecol Prog Ser 160: 291-293. 1997 Table 1. Madracis mirabilis. Mean percent predation per porites furcata in Belize (Littler et al. 1989). The colony at each depth (n = 3 for colony 2 at 13 m and n = 4 for pattern of predation observed in this study was also all other colonies at both depths). Mean percent predation reported by Yeudecker (1979) for P, damlcornis. Fur- was significantly h~gherat 13 m than at 20 m (Wilcoxon's signnd-ranks test T, = 2, n = 8, p < 0.01 17) ther, Llttler et al. (1989) suggested that the degree of fish predation is influenced by proximity to refugia for the corallivores and coral palatability. However, these Colony Mean percent predation Mean percent predation at 13 m at 20 m factors do not appear to be applicable to Conch Reef where topographic complexity and refugia increase 1 61 0 with depth (pers. obs.) in opposition to the observed 2 7 28 3 125 21 increase in predation across the experimental depth, 4 108 26 and where deep water M. mirabilis is susceptible to 5 102 29 predation when transplanted to shallower depths. 6 75 17 7 37 32 Field studies examining the direct role of corallivo- 8 83 10 rous fishes on coral zonation are limited (Kaufman Mean of means 74.75 20.38 1977, Neudecker 1979, Wellington 198213, Littler et SE 12.90 3.61 al. 1989, reviewed by Glynn 1990, Hixon 1997). Our observations provide further evidence that fish preda- tion can be an important factor limiting the vertical condition (Fig. 1). These results indicate that at Conch distribution of reef corals. Reef predation by corallivorous fish plays a critical role in restricting the distribution of Madracis mirabilis to Acknowledgements. We thank the staff of the National depths below 13 m. Although we did not directly Undersea Research Center's Florida Keys Program (UNCW) observe fish predation on M. mirabilis since caged con- for their support during 'Aquarius' mission No. 17 and the trols were not employed in this experimental design, Jason Foundation for Education (Jason Project V11) for the cusp-like bite marks revealed that parrotfish and/ financial assistance. For their assistance in the fleld we thank or pufferfish are likely to be responsible. No other evi- K. Sebens, G. Taylor, K. Wood and D. J. Roller. We also thank Dr C. Birkeland and 3 anonymous reviewers for their dence of damage due to handling or predation by other helpful comments on the manuscript. organisms was observed. Situations where fish predation limits the zonation LITERATURE CITED patterns of corals have also been reported in the branching coral Pocillopora damicornis (Neudecker Brock RE (1979) An experimental study of the effects of 1979, Wellington 1982b) and massive Pavona spp. grazing by parrotfishes and role of refuges in benthic community structure. Mar Biol 51 :381-388 (Wellington 1982b) in the Pacific, for Montastraea Fishelson L (1973) Ecological and biological phenomena In- annularis and Acropora cervicornis in Jamaica (Kauf- fluencing coral-species composition on the reef tables at man 1977) and for Porites astreoides and Porites Eilat (Gulf of Aqaba, Red Sea). Mar Biol 19:183-196 Flg 1 Madraas mlrab~l~sfragments from 13 m w~thbranch tlps bltten off (on left) and from 20 m wlth lntact branch tlps (on nght) Grottoli-Everett & Wellington: Flsh predation on Madrac~sm~rabilis 293 Glynn PLV (1976) Some physlcal and biological determinants Littler MM Taylor PR. Llttler DS (1989) Complex lnteractlons of coral commun~tystructure In the Eastern Paciflc Ecol In the contlol of coral zonatlon on a Car~bbeanreef flat Monogr 46:431-456 Oecologia 80 331-340 Glynn PW (1988) El N~no-Southern Oscillation 1982-83: Loya Y (1972) Community structure and specles dlverslty of nearshore population, community and ecosystem re- hermatyp~ccorals at Ellat Red Sea Mar B101 13 100-123 sponses Annu Rev Ecol Syst 19 309-345 McClanahan TR, Kamukuru AT Muthiga NA Gllagabher Glynn PM' (1990) Feeding ecology of selected coral-reef Yebio M Obura D (1996) Effect of sea uichin leductions macroconsumers; patterns and effects on coral community on algae, coral and flsh populatlons Conserv Biol 10 structure. In: Dubinsky Z (ed) Ecosystems of the world 25. 136-154 Coral reefs. Elsevier, Amsterdam, p 365-400 McClanahan TR Shaf~rSH (1990) Causes and consequences Glynn PW, Colgan MW (1988) Sporadic disturbances in fluc- of sea ulchln abundance and dlvers~tyIn Kenyan coral tuating coral reef environments: El Nilio and coral reef reef lagoons Oecologla 83 362-370 development in the Eastern Paclf~c.Am Zool 32:707-718 Neudecker S (1979)Effects of grazlng and browsing flshes on Glynn PW, Stewart RH, McCosker JE (1972) Paclflc coral the zonatlon of corals in Guam Ecology 60 666-672 reefs of Panama: structure, dlstributlon and predators. Porter JW (1972) Patterns of species dlverslty In Canbbean Geol Rundsch 61:483-519 reef corals Ecology 53 745-748 Glynn PW, Wellington GM, Birkeland C (1979) Coral reef Porter JW (1974) Commun~tystructure of coral reefs on oppo- growth in the Galapagos: limitation by sea urchins. SCI- site sldes of the Isthmus of Panama Sclence 186 543-545 ence 203.47-49 Porter JW (1976) Autotrophy heterotrophy and resource par- Harmelln-Vivien ML, Bouchon-Navaro Y (1983) Feeding dlets tltloning In Caribbean reef-building corals Am Nat 110 and significance of coral feed~nganlong Chaetodontid 731-742 f~shesin Moorea (French Polynesia). Coral Reefs 2: Robertson R (1970) Review of the predators and parasites of 119-127 stony corals, with special reference to symbiotic proso- Hixon MA (1997) Effects of reef flshes on corals and algae In: branch gastropods Pac Sci 24 43-54 Birkeland C (ed) Life and death of coral reefs Chapman Sammarco PW (1980) Djadema and its relationsh~pto coral and Hall, New York, p 231-246 spat mortality grazlng competition and biological dlstur- Humann P (1993) Reef coral ~dentiflcatlon.New World Publi- bance J Exp Mar Blol Ecol45 245-272 cat~ons,Inc, Jacksonville Sammarco PW (1982) Echlnoid grazing as a structunng force Huston MA (1985) Patterns of species diversity on coral reefs.
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