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Selection by Two Decapod Crabs (<I>Percnon Gibbesi</I> and <I>Stenorhynchus Seticornis</I>)

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Selection by Two Decapod Crabs (<I>Percnon Gibbesi</I> and <I>Stenorhynchus Seticornis</I>) BULLETIN OF MARINE SCIENCE, 63(1): 241–247, 1998 CORAL REEF PAPER SELECTION BY TWO DECAPOD CRABS (PERCNON GIBBESI AND STENORHYNCHUS SETICORNIS) ASSOCIATING WITH AN URCHIN (DIADEMA ANTILLARUM) AT TOBAGO, WEST INDIES Floyd E. Hayes, Victor L. Joseph, Horace S. Gurley and Brian Y. Y. Wong ABSTRACT We studied the relationships among two species of decapod crabs, Percnon gibbesi and Stenorhynchus seticornis, associating with an urchin, Diadema antillarum, in shallow water along the coasts of Tobago, West Indies. The crabs did not associate with other shorter-spined urchin species, suggesting that the crabs gained protection by associating with the relatively long-spined D. antillarum. Most D. antillarum (83.4%; n = 259) lacked an associating crab; P. gibbesi associated with D. antillarum more often (11.6% of ur- chins) than did S. seticornis (3.5%) and an unidentified crab (1.5%). We found no evi- dence that P. gibbesi and S. seticornis differed in their selection of D. antillarum hosts. We also found no evidence that the crabs selected D. antillarum hosts providing a poten- tially safer microhabitat by having longer spines, occurring in denser populations, occur- ring nearer to each other, or occurring in deeper water (within the depth range of this study). These results suggest that selection of D. antillarum hosts by P. gibbesi and S. seticornis is random, with minimal interspecific competition. Decapod crustaceans form symbiotic relationships with a variety of hosts, including echinoderms (Ross, 1983). However, the ecological relationships of most symbioses re- main poorly studied. In the Caribbean Sea, the spines of the long-spined urchin, D. antillarum, have been reported to provide refuge for a variety of commensal organisms, including an anemone, a flatworm, a copepod, a mysid shrimp, the young of many fish species, and a variety of decapod crustaceans including several palaemonid shrimp (Periclimenes sp., Stegopontonia sp., Tuleariocaris neglecta), a spiny lobster (Panulirus argus), and a grapsid crab (Percnon gibbesi; Randall et al., 1964; Chase, 1969; Clifton et al., 1970; Davis, 1971; Castro, 1974; Gooding, 1974; Serafy, 1979; Criales, 1984). The urchin crab, P. gibbesi, is a well known associate of D. antillarum (Schmalfuss, 1976; Adey et al., 1977; Powers, 1977; Colin, 1978; Kaplan, 1988; Humann, 1992). Colin (1978) stated that the crab gains protection from the relatively long, mildly toxic spines of the urchin, but the urchin apparently derives no benefit from the crab. The arrow crab, Stenorhynchus seticornis, has been observed closely associating with sponges, stony cor- als, soft corals, gorgonians, anemones and crinoids in the Bahamas and Virgin Islands (Barr, 1971, 1975; Herrnkind et al., 1976; Stanton, 1977; Schriever, 1978). However, there are only a few anecdotal reports of S. seticornis associating with urchins. Sefton and Webster (1986) reported that S. seticornis associated with urchins, but did not specify which species. When attempting to capture individual S. seticornis, Barr (1971: 220; 1975: 55) noted that “At times the crab would retreat among the spines of one or more urchins (Diadema). Whether this response was by design or chance is unknown, but the net result was extremely effective.” Adey et al. (1977: 36 and 49) stated that “Percnon gibbesi and Stenorhynchus hispidus are common near Diadema, crevices and sometimes anemones” in the Virgin Islands. However, it remains uncertain whether Adey et al. (1977) were referring to S. seticornis, of which S. hispidus is not a known synonym (Goeke, 1989), or the coral spider crab, Mithrax hispidus. 241 242 BULLETIN OF MARINE SCIENCE, VOL. 63, NO. 1, 1998 During excursions in June and November 1994 to Tobago, we repeatedly observed both P. gibbesi and S. seticornis within 5 cm of and retreating under the spines of D. antillarum in shallow water (<5 m deep) along the coasts. We subsequently returned to Tobago to further study the relationships among these species and a third unidentified crab associat- ing with D. antillarum. This study reports for the first time the association of S. seticornis with D. antillarum. We tested the hypothesis that the three species of crabs differed in their selection of urchin hosts as a consequence of interspecific competition. We also tested the hypothesis that the crabs selected D. antillarum hosts providing a potentially safer microhabitat by (1) having longer spines, (2) occurring in greater densities, (3) occurring nearer to each other and (4) occurring in deeper water. STUDY AREA AND METHODS The island of Tobago is perched along the edge of the South American continental shelf in the southeastern Caribbean Sea. The marine environment of Tobago is dominated by the northward flowing Guyana current, and is seasonally influenced by discharges of freshwater and sediments from South American rivers (Kenny and Bacon, 1981). The coasts of Tobago are characterized by steeply sloping volcanic rock formations alternating with sandy embayments, with the rock forma- tions forming a foundation for patches of fringing coral reefs (Laydoo, 1985, 1991). More exten- sive fringing reefs occur on an elevated limestone platform along the southwestern coast of Tobago (Kenny, 1976; Kenny and Bacon, 1981; Laydoo, 1985, 1991). The coral reef ecosystems of Tobago have been described by Kenny (1976), Ramsaroop (1981) and Laydoo (1985, 1991). The maximum tidal range is about 1.3 m (Kenny and Bacon, 1981). During 22–24 March 1995 and 5 September 1996, we carefully examined 259 D. antillarum urchins during daylight on rock or coral substrates in shallow water (<5 m deep) at Mount Irvine Bay, Arnos Vale Bay, Bloody Bay and Man-of-War Bay; each locality is situated along the northern coast of Tobago. We recorded the species of crab and the number of individuals present within 5 cm of each D. antillarum. Four specimens of S. seticornis were later collected from the urchins and three from nearby gorgonians at Mount Irvine Bay; each was examined under a dissecting micro- scope and distinguished from the sympatric but deeper water (>30 m, thus unlikely to occur) S. yangi by the presence of dense felt and setae on the rostrum (Goeke, 1989). For each D. antillarum examined, we measured the following variables: (1) length (cm) of the longest spines, at the top of the urchin (using a transparent plastic ruler inserted between spines); (2) number of conspecific individuals within a 0.57 m radius (=1.0 m2); (3) number of individuals of other urchin species (an estimated 98% rock-boring urchins, Echinometra lucunter, and 2% slate-pencil urchins, Eucidaris tribuloides) within a 0.57 m radius (=1.0 m2); (4) distance (m) to the nearest conspecific individual; and (5) depth (estimated to nearest 0.5m) below the surface. Because of the low frequency of crabs associating with D. antillarum, the data from all four localities were pooled for subsequent analyses. A one-sample chi-square test (Χ2 statistic; Zar, 1984) was used to compare the proportions of crab species associating with D. antillarum. A one- way anova (F statistic; Zar, 1984) was used to test for differences in spine length and depth below the surface among D. antillarum with or without crabs; a posteriori multiple comparison tests were computed using student’s t test (t statistic; Zar 1984) with Bonferroni adjustments of the alpha level (Lentner and Bishop, 1986). Kruskal-Wallis tests (H statistic; Zar, 1984) were used to test for differences in density and nearest conspecific distance among D. antillarum with or without crabs. A student’s t test was used to compare spine length between D. antillarum with either one or two P. gibbesi. All tests were computed using Statistix 3.1 software (Anonymous, 1990). HAYES ET AL.: CRAB-URCHIN ASSOCIATIONS 243 ybnoitcelestsohgnitceffayllaitnetopselbairaV.1elbaT isebbignoncreP , suhcnyhronetS sinrocites nihcruehthtiwgnitaicossabarcdeifitnedinunadna .murallitnaamedaiD INelbairavtnednepedn MDEASNMXIMNA Snipeelnhtgc(m)foD.aurallitnma noarcbeserptn 10.24208.560.9.1621 P.gibbesip8resent 171.720.965.7.103 S.seticornisp6resent 110.520.080.5.19 u5nidentifiedcrabpresent 80.220.550.1.14 Density(within0.57mradius)ofD.antillarumb n5ocrabpresent 15.71162.7 20 .Pbigisebp0tneser 29.01168.3 2 sinrocites.S p7tneser 13.61136.0 u5deifitnedin barc tneserp 10.20124.5 Dytisne(wnihti5.07m)suidarfollanihcru seicepsc n6obarc tneserp 23.4216.2 1220 P.gibbesip5resent 31.7416.3 182 S.seticornisp7resent 32.1311.9 16 u5nidentifiedcrabpresent 10.20124.5 Decnatsi(mot)tseraen.Daurallitnmd n4obarc tneserp 19.322.300.0.02220 P.gibbesip8resent 28.141.500.0.282 S.seticornisp0resent 25.731.800.0.16 u8nidentifiedcrabpresent 19.721.100..54 Depth(m)ofD.antillarumfromsurfacee n8ocrabpresent 29.005.805..4621 P.gibbesip0resent 23.005.805..303 S.seticornisp1resent 27.615.115..49 u0nidentifiedcrabpresent 12.505.805..24 aF=3.06,P=0.028;bH=1.83,P=0.61;cH=3.30,P=0.35;dH=0.82,P=0.84;eF=1.72,P=0.16 RESULTS In the shallow waters of Tobago, P. gibbesi was ubiquitous on rocks both above and below the surface of the ocean within or just below the intertidal zone, and often did not associate with D. antillarum. In contrast, S. seticornis was considerably less common than P. gibbesi, associating almost exclusively with D. antillarum and an unidentified gorgonian at a minimum depth of 1 m. An unidentified red crab (probably Mithrax sp.) that eluded capture was found associating only with D. antillarum in shallow water. When found on the substrate within 5 cm of D. antillarum, each crab species quickly retreated under the urchin’s spines. The crabs followed the urchin when it moved as we measured and examined the urchin for associating crabs. Of 259 D. antillarum examined, 43 (16.6%) had an associated crab; P. gibbesi was present with 30 urchins (11.6%), S. seticornis was present with nine urchins (3.5%), and an unidentified crab was present with four urchins (1.5%). Of the 30 D. antillarum host- ing P.
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