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Responses of Two Coral Reef Toadfishes (Batrachoididae) to the Demise of Their Primary Prey, the Sea Urchin Diadema Antillarum Author(S): D

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Responses of Two Coral Reef Toadfishes (Batrachoididae) to the Demise of Their Primary Prey, the Sea Urchin Diadema Antillarum Author(S): D Responses of Two Coral Reef Toadfishes (Batrachoididae) to the Demise of Their Primary Prey, the Sea Urchin Diadema antillarum Author(s): D. Ross Robertson Reviewed work(s): Source: Copeia, Vol. 1987, No. 3 (Aug. 5, 1987), pp. 637-642 Published by: American Society of Ichthyologists and Herpetologists (ASIH) Stable URL: http://www.jstor.org/stable/1445655 . Accessed: 27/11/2012 15:23 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Society of Ichthyologists and Herpetologists (ASIH) is collaborating with JSTOR to digitize, preserve and extend access to Copeia. http://www.jstor.org This content downloaded by the authorized user from 192.168.72.223 on Tue, 27 Nov 2012 15:23:23 PM All use subject to JSTOR Terms and Conditions MATTHEWS-VARIATION IN CYPRINELLA 637 American minnows, with emphasis on the genus , ANDN. L. ODEN. 1978a. Spatial autocorre- Cyprinella(Teleostei: Cypriniformes).Misc. Publ. lation in biology. I. Methodology. Biol. J. Linnean Univ. KansasMus. Nat. Hist. In press. Soc. 10:199-228. METCALF,A. L. 1966. Fishes of the Kansas River ,AND . 1978b. Spatial autocorrelation system in relation to zoogeography of the Great in biology. II. Some biologicalimplications and four Plains. Univ. Kans. Publ. Mus. Nat. Hist. 17:23- applicationsof evolutionaryand ecological interest. 189. Ibid. 10:229-249. PAGE,L. M., AND R. L. SMITH. 1970. Recent range SOULE,M. 1973. The epistasis cycle: a theory of mar- adjustments and hybridization of Notropis lutrensis ginal populations. Ann. Rev. Ecol. Syst. 4:165-188. and Notropisspilopterus in Illinois. Trans. Ill. State UNDERHILL,J. C. 1960. Variation in the red shiner, Acad. Sci. 63:264-272. Notropis lutrensis (Baird and Girard). Minnesota ROBINS, C. R., R. M. BAILEY, C. E. BOND,J. R. BROOK- Acad. Sci. 28:78-80. ER, E. A. LACHNER,R. N. LEA AND W. B. SCOTT. WOOTEN,M. C. 1983. Genetic and morphological 1980. A list of common and scientific names of variation in natural populations of the red shiner, fishes from the United States and Canada (4th ed.). Notropis lutrensis and their relationship to adapta- Amer. Fish. Soc. Spec. Publ. No. 12. tion in a generalist species. Unpubl. Ph.D. dissert., SMITH,P. W. 1979. The fishes of Illinois. University North Texas State University, Denton, Texas. of Illinois Press, Urbana, Illinois. SOKAL,R. R. 1979. Testing statistical significance of BIOLOGICAL STATION AND DEPARTMENT OF geographic variation patterns. Syst. Zool. 28:227- ZOOLOGY, UNIVERSITY OF OKLAHOMA, 232. KINGSTON,OKLAHOMA 73439. 27 . 1983. character variation in Accepted Analyzing geo- Nov. 1986. graphic space., p. 384-403. In: Numerical taxon- omy. J. Felsenstein (ed.). Springer-Verlag, New York, New York. Copeia, 1987(3), pp. 637-642 Responses of Two Coral Reef Toadfishes (Batrachoididae) to the Demise of Their Primary Prey, the Sea Urchin Diadema antillarum D. Ross ROBERTSON In 1983-84 the sea urchin Diadema antillarum, which was abundant on coral reefs throughout the Caribbean, suffered a mass mortality. Its densities decreased by about 95% throughout most of its range. In Panama, prior to that mortality event, two reef toadfishes, Amphichthys cryptocentrus and Sanopus barbatus fed almost exclusively on Diadema. Two and a half years after that event the abun- dance of at least one toadfish appears to be at the pre-event level and both species are actively breeding. After the event these "specialist" fishes changed their diets in different ways. The diet of A. cryptocentrus has become generalized and now includes a broad range of mobile benthic invertebrates (crabs, hermit crabs, gastropods, octopods, echinoids). The diet of S. barbatus now consists primarily of fishes, but also includes a few mobile benthic invertebrates. The ability of Diadema predators to maintain their populations at high levels may reduce the potential for Diadema populations to recover. U NTIL recently the sea urchin Diadema an- Caribbean (Randall, 1967). In Jan.-May 1983 tillarum was an abundant and ecologically populations of Diadema on the Caribbean coast influential organism on Caribbean coral reefs of Panama suffered a rapid mass mortality, which (see references cited in Lessios et al., 1984a, subsequently affected the species throughout 1984b). Many reef fishes included this urchin most of its geographic range (Lessios, et al., in their diets and it constituted the major item 1984a, 1984b; Hughes et al., 1985). In San Bias eaten by several species at sites in the eastern the mortality reduced Diadema populations by ? 1987 by the American Society of Ichthyologists and Herpetologists This content downloaded by the authorized user from 192.168.72.223 on Tue, 27 Nov 2012 15:23:23 PM All use subject to JSTOR Terms and Conditions 638 COPEIA, 1987, NO. 3 - 1Km Fig. 1. Patch reefs of San Bias Point. Solid lines define emergent reefs, dashed lines define submerged reefs. Substratebetween reefs consists of sand, mainly covered with macroalgaeand seagrasses. 95% (Lessios et al., 1984a) and those popula- Hoffman and Robertson (1983) found that both tions did not recover in the subsequent 3 yr (H. species were most likely to have fresh material A. Lessios, pers. comm.). in their guts in the morning. Consequently the Prior to the mortality event two species of 1984-86 series of specimens were collected in San Bias toadfishes (Batrachoididae), Amphi- the morning. Fishes were collected with Quin- chthyscryptocentrus and Sanopus barbatus, fed al- aldine anaesthetic and dissected within 1 h of most exclusively on D. antillarum (Hoffman and capture. All identifiable material in the guts was Robertson, 1983). Here I describe the re- recorded and measured. sponses of those fishes to the abrupt loss of their primary prey. Reproductiveactivity. -The degree of activity of the ovaries of females collected for gut analyses METHODS was recorded. Ripe ovaries are readily recog- nizable since both species produce eggs about Study area.-Data were collected at the same 5 mm in diameter. Hoffman and Robertson site in the Archipielago de San Bias (Fig. 1) both (1983) found males of A. cryptocentruswith ag- before (Hoffman and Robertson, 1983) and af- gregations of small juveniles (2-3 cm long) in ter the Diadema mortality. their burrows and searches were made for such juveniles in the shelters of both species during Gut contentanalyses.-The mortality affected San the 1984-86 series of collections. Bias populations of D. antillarum in April-May 1983 (Lessios et al., 1984a, 1984b). Information Population change in A. cryptocentrus.- on the toadfishes' diets was collected during Amphichthyscryptocentrus was by far the more 1977 and 1981 (Hoffman and Robertson [1983]) abundant of the two toadfishes before the Dia- and Sept. 1983-Jan. 1986. Specimens were col- dema mortality (Hoffman and Robertson, 1983). lected from shallow reefs immediately to the During a 6 mo period in 1978 a series of cen- east of Punta de San Bias (Fig. 1) in both cases. suses was made of the A. cryptocentruspopulation This content downloaded by the authorized user from 192.168.72.223 on Tue, 27 Nov 2012 15:23:23 PM All use subject to JSTOR Terms and Conditions ROBERTSON-DEMISE OF TOADFISH PREY 639 TABLE 1. GUT CONTENTS OF Amphichthyscryptocentrus BEFORE AND AFTER Diadema MORTALITY. Before" After Proportion of Proportion of Proportion of items in total Proportion of items in total Sizeb of items Item fish with items gut contents fish with items gut contents (cm) Echinoids Diadema .80 .85 .03 .01 8.0 Echinometraand Eucidaris .00 0 .20 .06 4.0 (3.0-5.0) Crabs .06 .08 .66 .36 1.8 (1.0-6.0) Gastropods .06 .08 .40 .32 1.1 (0.5-4.5) Hermit crabs .00 0 .29 .15 1.6 (0.8-5.0) Octopods .00 0 .11 .05 20.0 (15.0-36.0) Scallops .00 0 .06 .02 2.1 (2.0-2.2) Lobster .00 0 .03 .01 3.0 Fishes .00 0 .03 .01 4.0 Empty .09 .09 N 35 39 35 94 Data from Hoffman and Robertson (1983). b X (range). Echinoids = test diameter, crabs = carapace width, gastropods and hermit crabs = shell length, octopods = arm tip to tip, lobster/ shrimp = head to tail length, fish = total length, scallop = shell width. in one 0.225 ha area immediately to the west cryptocentrus(Table 1). In terms of numerical of Smithsoniantupo (Fig.l). Since May 1983, abundance, the main items present in A. cryp- when the Diadema mortality was in progress, tocentrusafter the mortality were crabs, gastro- this same area has been censused at approxi- pods, and hermit crabs. However, when prey mately monthly intervals. No estimate was made size also is taken into account, a larger variety of the population density of S. barbartus before of prey contributed substantially to the biomass or after the mortality event. of the diet of that toadfish. Individuals of the three most abundant prey groups (gastropods, hermit crabs, and crabs) typically were consid- RESULTS erably smaller than individuals of two less abun- dant prey groups (octopods and echinoids) (Ta- Gut analyses.-Amphichthys cryptocentrus.- ble 1). Thus all five of these prey types are Prior to the Diadema die-off that urchin rep- important components of the new diet of A. resented almost the entire gut contents of A. cryptocentrus. TABLE 2. GUT CONTENTS OF Sanopus barbatus BEFORE AND AFTER THE DiademaMORTALITY. Before' After Proportion of Proportion of Proportion of items in total Proportion of items in total Sizeb of items Item fish with items gut contents fish with items gut contents (cm) Diadema .92 1.00 .00 .00 Fishes .00 .00 .44 .48 12.0 (10.0-15.0) Shrimps .00 .00 .11 .10 3.5 (2.0-5.0) Scallops .00 .00 .06 .05 1.5 Hermit crabs .00 .00 .11 .29 1.7 (1.2-2.5) Crabs .00 .00 .06 .05 2.0 Gastropods .00 .00 .06 .05 1.0 Empty .08 - .28 N 13 13 18 21 Data from Hoffman and Robertson (1983).
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