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Predation, Prey Refuges, and the Structure of Coral-Reef Fish Assemblages

We studied the influence of piscivorous fishes and prey refuges on assemblages of fishes occupying 52 model reefs in a large seagrass bed off St. Thomas, U.S. Virgin Islands. We conducted three experiments: two involving 6 reefs each, lasting 2 and 5 yr, and one involving 40 reefs, lasting 1 yr. Eac...

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Published in:Ecological monographs 1993-02, Vol.63 (1), p.77-101
Main Authors: Hixon, Mark A., Beets, James P.
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Beets, James P.
description We studied the influence of piscivorous fishes and prey refuges on assemblages of fishes occupying 52 model reefs in a large seagrass bed off St. Thomas, U.S. Virgin Islands. We conducted three experiments: two involving 6 reefs each, lasting 2 and 5 yr, and one involving 40 reefs, lasting 1 yr. Each experiment included replicate reefs in various combinations of five structural treatments: holeless controls, 12 and 24 small holes, and 12 and 24 large holes. Tagging studies indicated that the reefs were sufficiently isolated from each other to comprise statistically independent replicates, and that resident piscivores occupied home reefs. We observed 97 species on or near the reefs, representing all major foraging guilds, and each holed reef supported hundreds of individuals. We examined four categories of fish: (1) large reef associates (too large for the small holes; most of these fish were both predators on smaller fish and prey for larger transient piscivores), (2) moray eels (piscivores that could fit into the small holes), (3) small reef associates (potential prey that could fit into the small holes), and (4) juvenile grunts (potential prey that sporadically were extremely abundant). We tested five a priori predictions of the general hypothesis that predation is an important process structuring reef-fish assemblages. The first two predictions dealt with the role of prey refuges. First, if reef holes function as prey refuges, then prey fish should be most abundant on reefs providing holes near their body diameters, because such holes would make the prey fish safest from predation. Seven of eight experimental comparisons supported this prediction, and five of them were statistically significant. Second, if refuge availability limits prey abundance, then prey fish should be more abundant on reefs with 12 holes than those with no holes, and should be more abundant on reefs with 24 holes than those with 12 holes. The first part of this prediction was verified by all nine experimental comparisons, seven of which were statistically significant. However, there were no strong differences between 12-hole and 24-hole reefs. Thus, between 0 and 12 holes per reef, holes limited local prey populations; between 12 and 24 holes per reef, the number of holes was not limiting. Several lines of evidence suggested that the latter pattern was due to temporary saturation of the study area with refuges when we added 40 reefs to 12 existing reefs. The remaining three predict
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We conducted three experiments: two involving 6 reefs each, lasting 2 and 5 yr, and one involving 40 reefs, lasting 1 yr. Each experiment included replicate reefs in various combinations of five structural treatments: holeless controls, 12 and 24 small holes, and 12 and 24 large holes. Tagging studies indicated that the reefs were sufficiently isolated from each other to comprise statistically independent replicates, and that resident piscivores occupied home reefs. We observed 97 species on or near the reefs, representing all major foraging guilds, and each holed reef supported hundreds of individuals. We examined four categories of fish: (1) large reef associates (too large for the small holes; most of these fish were both predators on smaller fish and prey for larger transient piscivores), (2) moray eels (piscivores that could fit into the small holes), (3) small reef associates (potential prey that could fit into the small holes), and (4) juvenile grunts (potential prey that sporadically were extremely abundant). We tested five a priori predictions of the general hypothesis that predation is an important process structuring reef-fish assemblages. The first two predictions dealt with the role of prey refuges. First, if reef holes function as prey refuges, then prey fish should be most abundant on reefs providing holes near their body diameters, because such holes would make the prey fish safest from predation. Seven of eight experimental comparisons supported this prediction, and five of them were statistically significant. Second, if refuge availability limits prey abundance, then prey fish should be more abundant on reefs with 12 holes than those with no holes, and should be more abundant on reefs with 24 holes than those with 12 holes. The first part of this prediction was verified by all nine experimental comparisons, seven of which were statistically significant. However, there were no strong differences between 12-hole and 24-hole reefs. Thus, between 0 and 12 holes per reef, holes limited local prey populations; between 12 and 24 holes per reef, the number of holes was not limiting. Several lines of evidence suggested that the latter pattern was due to temporary saturation of the study area with refuges when we added 40 reefs to 12 existing reefs. The remaining three predictions dealt directly with the community-level role of predation. First, predators should affect local prey abundance either chronically, in which case a negative relationship among reefs is predicted between the average abundances of predators and prey, or sporadically, in which case a negative relationship is predicted between the abundance of predators and the maximum number of co-occurring prey ever observed at each predator abundance. The former prediction was falsified, whereas the latter was verified. Observations of extreme type III survivorship of recruit cohorts on reefs with many piscivores and occasional direct observations of piscivory bolstered the conclusion that this relationship was causal. Finally, we predicted that predators should affect the number of prey species on a reef. We observed a significant negative relationship among reefs between predator abundance and maximum prey-species richness. Comparing species' relative abundances on reefs at the extremes of this regression, piscivores appear to have nonselectively reduced and extirpated both common and rare prey species, although this relationship remains purely correlative. In our model system, high local species diversity appears to have been maintained despite rather than because of predation. We propose a conceptual model where the local abundances of coral-reef fishes are determined by the relative magnitudes of recruitment by larvae, colonization by juveniles and adults, predation, and competition for refuges, each of which varies through time and space. 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Psychology ; Marine ; Marine biology ; Marine ecology ; Marine fishes ; Pisces ; Predation ; Predators ; Sea water ecosystems ; Synecology</subject><ispartof>Ecological monographs, 1993-02, Vol.63 (1), p.77-101</ispartof><rights>Copyright 1993 The Ecological Society of America</rights><rights>1993 by the Ecological Society of America</rights><rights>1993 INIST-CNRS</rights><rights>Copyright Ecological Society of America Feb 1993</rights><lds50>peer_reviewed</lds50><woscitedreferencessubscribed>false</woscitedreferencessubscribed><citedby>FETCH-LOGICAL-c4757-e61f3f9246a92fccbce113d450d9e56b6907fa091900b7dc891d739127602d5a3</citedby></display><links><openurl>$$Topenurl_article</openurl><openurlfulltext>$$Topenurlfull_article</openurlfulltext><thumbnail>$$Tsyndetics_thumb_exl</thumbnail><linktopdf>$$Uhttps://www.jstor.org/stable/pdf/2937124$$EPDF$$P50$$Gjstor$$H</linktopdf><linktohtml>$$Uhttps://www.jstor.org/stable/2937124$$EHTML$$P50$$Gjstor$$H</linktohtml><link.rule.ids>314,780,784,27924,27925,58238,58471</link.rule.ids><backlink>$$Uhttp://pascal-francis.inist.fr/vibad/index.php?action=getRecordDetail&amp;idt=4649061$$DView record in Pascal Francis$$Hfree_for_read</backlink></links><search><creatorcontrib>Hixon, Mark A.</creatorcontrib><creatorcontrib>Beets, James P.</creatorcontrib><title>Predation, Prey Refuges, and the Structure of Coral-Reef Fish Assemblages</title><title>Ecological monographs</title><description>We studied the influence of piscivorous fishes and prey refuges on assemblages of fishes occupying 52 model reefs in a large seagrass bed off St. Thomas, U.S. Virgin Islands. We conducted three experiments: two involving 6 reefs each, lasting 2 and 5 yr, and one involving 40 reefs, lasting 1 yr. Each experiment included replicate reefs in various combinations of five structural treatments: holeless controls, 12 and 24 small holes, and 12 and 24 large holes. Tagging studies indicated that the reefs were sufficiently isolated from each other to comprise statistically independent replicates, and that resident piscivores occupied home reefs. We observed 97 species on or near the reefs, representing all major foraging guilds, and each holed reef supported hundreds of individuals. We examined four categories of fish: (1) large reef associates (too large for the small holes; most of these fish were both predators on smaller fish and prey for larger transient piscivores), (2) moray eels (piscivores that could fit into the small holes), (3) small reef associates (potential prey that could fit into the small holes), and (4) juvenile grunts (potential prey that sporadically were extremely abundant). We tested five a priori predictions of the general hypothesis that predation is an important process structuring reef-fish assemblages. The first two predictions dealt with the role of prey refuges. First, if reef holes function as prey refuges, then prey fish should be most abundant on reefs providing holes near their body diameters, because such holes would make the prey fish safest from predation. Seven of eight experimental comparisons supported this prediction, and five of them were statistically significant. Second, if refuge availability limits prey abundance, then prey fish should be more abundant on reefs with 12 holes than those with no holes, and should be more abundant on reefs with 24 holes than those with 12 holes. The first part of this prediction was verified by all nine experimental comparisons, seven of which were statistically significant. However, there were no strong differences between 12-hole and 24-hole reefs. Thus, between 0 and 12 holes per reef, holes limited local prey populations; between 12 and 24 holes per reef, the number of holes was not limiting. Several lines of evidence suggested that the latter pattern was due to temporary saturation of the study area with refuges when we added 40 reefs to 12 existing reefs. The remaining three predictions dealt directly with the community-level role of predation. First, predators should affect local prey abundance either chronically, in which case a negative relationship among reefs is predicted between the average abundances of predators and prey, or sporadically, in which case a negative relationship is predicted between the abundance of predators and the maximum number of co-occurring prey ever observed at each predator abundance. The former prediction was falsified, whereas the latter was verified. Observations of extreme type III survivorship of recruit cohorts on reefs with many piscivores and occasional direct observations of piscivory bolstered the conclusion that this relationship was causal. Finally, we predicted that predators should affect the number of prey species on a reef. We observed a significant negative relationship among reefs between predator abundance and maximum prey-species richness. Comparing species' relative abundances on reefs at the extremes of this regression, piscivores appear to have nonselectively reduced and extirpated both common and rare prey species, although this relationship remains purely correlative. In our model system, high local species diversity appears to have been maintained despite rather than because of predation. We propose a conceptual model where the local abundances of coral-reef fishes are determined by the relative magnitudes of recruitment by larvae, colonization by juveniles and adults, predation, and competition for refuges, each of which varies through time and space. 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Psychology</subject><subject>Marine</subject><subject>Marine biology</subject><subject>Marine ecology</subject><subject>Marine fishes</subject><subject>Pisces</subject><subject>Predation</subject><subject>Predators</subject><subject>Sea water ecosystems</subject><subject>Synecology</subject><issn>0012-9615</issn><issn>1557-7015</issn><fulltext>true</fulltext><rsrctype>article</rsrctype><creationdate>1993</creationdate><recordtype>article</recordtype><recordid>eNp1kFFrFDEQx4MoeFbxKwSV-tLVmSSbOI_laLVQUdr6vOSyE7vH3m5NdpH79ua4w0LhnmYGfvPjPyPEW4RPSoP7rEg7VOaZWGBdu8oB1s_FAgBVRRbrl-JVzmvYzUQLcfUzceunbhzOZGm38obj_JvzmfRDK6d7lrdTmsM0J5ZjlMsx-b66YY7yssv38jxn3qx6XzZeixfR95nfHOqJ-HV5cbf8Vl3_-Hq1PL-ugnElD1uMOpIy1pOKIawCI-rW1NAS13ZlCVz0QEgAK9eGL4St04TKWVBt7fWJON17H9L4Z-Y8NZsuB-57P_A45wat1eVoVcB3T8D1OKehZGuU1lgDEBTo_TEINYAxztBO9XFPhTTmnDg2D6nb-LRtEJrd15vD1wv54eDzOfg-Jj-ELv_HjTUEFguGe-xv1_P2mK25WH5HIm01OveoXudpTEcT_AMsIpYa</recordid><startdate>199302</startdate><enddate>199302</enddate><creator>Hixon, Mark A.</creator><creator>Beets, James P.</creator><general>The Ecological Society of America</general><general>Ecological Society of America</general><general>Duke University Press</general><scope>IQODW</scope><scope>AAYXX</scope><scope>CITATION</scope><scope>HFXKP</scope><scope>IZSXY</scope><scope>K30</scope><scope>PAAUG</scope><scope>PAWHS</scope><scope>PAWZZ</scope><scope>PAXOH</scope><scope>PBHAV</scope><scope>PBQSW</scope><scope>PBYQZ</scope><scope>PCIWU</scope><scope>PCMID</scope><scope>PCZJX</scope><scope>PDGRG</scope><scope>PDWWI</scope><scope>PETMR</scope><scope>PFVGT</scope><scope>PGXDX</scope><scope>PIHIL</scope><scope>PISVA</scope><scope>PJCTQ</scope><scope>PJTMS</scope><scope>PLCHJ</scope><scope>PMHAD</scope><scope>PNQDJ</scope><scope>POUND</scope><scope>PPLAD</scope><scope>PQAPC</scope><scope>PQCAN</scope><scope>PQCMW</scope><scope>PQEME</scope><scope>PQHKH</scope><scope>PQMID</scope><scope>PQNCT</scope><scope>PQNET</scope><scope>PQSCT</scope><scope>PQSET</scope><scope>PSVJG</scope><scope>PVMQY</scope><scope>PZGFC</scope><scope>7QG</scope><scope>7SN</scope><scope>7SS</scope><scope>C1K</scope><scope>M7N</scope><scope>F1W</scope><scope>H95</scope><scope>L.G</scope></search><sort><creationdate>199302</creationdate><title>Predation, Prey Refuges, and the Structure of Coral-Reef Fish Assemblages</title><author>Hixon, Mark A. ; Beets, James P.</author></sort><facets><frbrtype>5</frbrtype><frbrgroupid>cdi_FETCH-LOGICAL-c4757-e61f3f9246a92fccbce113d450d9e56b6907fa091900b7dc891d739127602d5a3</frbrgroupid><rsrctype>articles</rsrctype><prefilter>articles</prefilter><language>eng</language><creationdate>1993</creationdate><topic>Animal and plant ecology</topic><topic>Animal, plant and microbial ecology</topic><topic>Biological and medical sciences</topic><topic>Coral reefs</topic><topic>Ecological competition</topic><topic>Ecology</topic><topic>Eels</topic><topic>Fish</topic><topic>Fundamental and applied biological sciences. Psychology</topic><topic>Marine</topic><topic>Marine biology</topic><topic>Marine ecology</topic><topic>Marine fishes</topic><topic>Pisces</topic><topic>Predation</topic><topic>Predators</topic><topic>Sea water ecosystems</topic><topic>Synecology</topic><toplevel>peer_reviewed</toplevel><toplevel>online_resources</toplevel><creatorcontrib>Hixon, Mark A.</creatorcontrib><creatorcontrib>Beets, James P.</creatorcontrib><collection>Pascal-Francis</collection><collection>CrossRef</collection><collection>Periodicals Index Online Segment 17</collection><collection>Periodicals Index Online Segment 30</collection><collection>Periodicals Index Online</collection><collection>Primary Sources Access—Foundation Edition (Plan E) - West</collection><collection>Primary Sources Access (Plan D) - International</collection><collection>Primary Sources Access &amp; Build (Plan A) - MEA</collection><collection>Primary Sources Access—Foundation Edition (Plan E) - Midwest</collection><collection>Primary Sources Access—Foundation Edition (Plan E) - Northeast</collection><collection>Primary Sources Access (Plan D) - Southeast</collection><collection>Primary Sources Access (Plan D) - North Central</collection><collection>Primary Sources Access—Foundation Edition (Plan E) - Southeast</collection><collection>Primary Sources Access (Plan D) - South Central</collection><collection>Primary Sources Access &amp; Build (Plan A) - UK / I</collection><collection>Primary Sources Access (Plan D) - Canada</collection><collection>Primary Sources Access (Plan D) - EMEALA</collection><collection>Primary Sources Access—Foundation Edition (Plan E) - North Central</collection><collection>Primary Sources Access—Foundation Edition (Plan E) - South Central</collection><collection>Primary Sources Access &amp; Build (Plan A) - International</collection><collection>Primary Sources Access—Foundation Edition (Plan E) - International</collection><collection>Primary Sources Access (Plan D) - West</collection><collection>Periodicals Index Online Segments 1-50</collection><collection>Primary Sources Access (Plan D) - APAC</collection><collection>Primary Sources Access (Plan D) - Midwest</collection><collection>Primary Sources Access (Plan D) - MEA</collection><collection>Primary Sources Access—Foundation Edition (Plan E) - Canada</collection><collection>Primary Sources Access—Foundation Edition (Plan E) - UK / I</collection><collection>Primary Sources Access—Foundation Edition (Plan E) - EMEALA</collection><collection>Primary Sources Access &amp; Build (Plan A) - APAC</collection><collection>Primary Sources Access &amp; Build (Plan A) - Canada</collection><collection>Primary Sources Access &amp; Build (Plan A) - West</collection><collection>Primary Sources Access &amp; Build (Plan A) - EMEALA</collection><collection>Primary Sources Access (Plan D) - Northeast</collection><collection>Primary Sources Access &amp; Build (Plan A) - Midwest</collection><collection>Primary Sources Access &amp; Build (Plan A) - North Central</collection><collection>Primary Sources Access &amp; Build (Plan A) - Northeast</collection><collection>Primary Sources Access &amp; Build (Plan A) - South Central</collection><collection>Primary Sources Access &amp; Build (Plan A) - Southeast</collection><collection>Primary Sources Access (Plan D) - UK / I</collection><collection>Primary Sources Access—Foundation Edition (Plan E) - APAC</collection><collection>Primary Sources Access—Foundation Edition (Plan E) - MEA</collection><collection>Animal Behavior Abstracts</collection><collection>Ecology Abstracts</collection><collection>Entomology Abstracts (Full archive)</collection><collection>Environmental Sciences and Pollution Management</collection><collection>Algology Mycology and Protozoology Abstracts (Microbiology C)</collection><collection>ASFA: Aquatic Sciences and Fisheries Abstracts</collection><collection>Aquatic Science &amp; Fisheries Abstracts (ASFA) 1: Biological Sciences &amp; Living Resources</collection><collection>Aquatic Science &amp; Fisheries Abstracts (ASFA) Professional</collection><jtitle>Ecological monographs</jtitle></facets><delivery><delcategory>Remote Search Resource</delcategory><fulltext>fulltext</fulltext></delivery><addata><au>Hixon, Mark A.</au><au>Beets, James P.</au><format>journal</format><genre>article</genre><ristype>JOUR</ristype><atitle>Predation, Prey Refuges, and the Structure of Coral-Reef Fish Assemblages</atitle><jtitle>Ecological monographs</jtitle><date>1993-02</date><risdate>1993</risdate><volume>63</volume><issue>1</issue><spage>77</spage><epage>101</epage><pages>77-101</pages><issn>0012-9615</issn><eissn>1557-7015</eissn><coden>ECMOAQ</coden><abstract>We studied the influence of piscivorous fishes and prey refuges on assemblages of fishes occupying 52 model reefs in a large seagrass bed off St. Thomas, U.S. Virgin Islands. We conducted three experiments: two involving 6 reefs each, lasting 2 and 5 yr, and one involving 40 reefs, lasting 1 yr. Each experiment included replicate reefs in various combinations of five structural treatments: holeless controls, 12 and 24 small holes, and 12 and 24 large holes. Tagging studies indicated that the reefs were sufficiently isolated from each other to comprise statistically independent replicates, and that resident piscivores occupied home reefs. We observed 97 species on or near the reefs, representing all major foraging guilds, and each holed reef supported hundreds of individuals. We examined four categories of fish: (1) large reef associates (too large for the small holes; most of these fish were both predators on smaller fish and prey for larger transient piscivores), (2) moray eels (piscivores that could fit into the small holes), (3) small reef associates (potential prey that could fit into the small holes), and (4) juvenile grunts (potential prey that sporadically were extremely abundant). We tested five a priori predictions of the general hypothesis that predation is an important process structuring reef-fish assemblages. The first two predictions dealt with the role of prey refuges. First, if reef holes function as prey refuges, then prey fish should be most abundant on reefs providing holes near their body diameters, because such holes would make the prey fish safest from predation. Seven of eight experimental comparisons supported this prediction, and five of them were statistically significant. Second, if refuge availability limits prey abundance, then prey fish should be more abundant on reefs with 12 holes than those with no holes, and should be more abundant on reefs with 24 holes than those with 12 holes. The first part of this prediction was verified by all nine experimental comparisons, seven of which were statistically significant. However, there were no strong differences between 12-hole and 24-hole reefs. Thus, between 0 and 12 holes per reef, holes limited local prey populations; between 12 and 24 holes per reef, the number of holes was not limiting. Several lines of evidence suggested that the latter pattern was due to temporary saturation of the study area with refuges when we added 40 reefs to 12 existing reefs. The remaining three predictions dealt directly with the community-level role of predation. First, predators should affect local prey abundance either chronically, in which case a negative relationship among reefs is predicted between the average abundances of predators and prey, or sporadically, in which case a negative relationship is predicted between the abundance of predators and the maximum number of co-occurring prey ever observed at each predator abundance. The former prediction was falsified, whereas the latter was verified. Observations of extreme type III survivorship of recruit cohorts on reefs with many piscivores and occasional direct observations of piscivory bolstered the conclusion that this relationship was causal. Finally, we predicted that predators should affect the number of prey species on a reef. We observed a significant negative relationship among reefs between predator abundance and maximum prey-species richness. Comparing species' relative abundances on reefs at the extremes of this regression, piscivores appear to have nonselectively reduced and extirpated both common and rare prey species, although this relationship remains purely correlative. In our model system, high local species diversity appears to have been maintained despite rather than because of predation. We propose a conceptual model where the local abundances of coral-reef fishes are determined by the relative magnitudes of recruitment by larvae, colonization by juveniles and adults, predation, and competition for refuges, each of which varies through time and space. Multifactorial field experiments will be necessary to test such pluralistic hypotheses.</abstract><cop>Washington, DC</cop><pub>The Ecological Society of America</pub><doi>10.2307/2937124</doi><tpages>25</tpages></addata></record>
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ispartof Ecological monographs, 1993-02, Vol.63 (1), p.77-101
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language eng
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source JSTOR Archival Journals and Primary Sources Collection
subjects Animal and plant ecology
Animal, plant and microbial ecology
Biological and medical sciences
Coral reefs
Ecological competition
Ecology
Eels
Fish
Fundamental and applied biological sciences. Psychology
Marine
Marine biology
Marine ecology
Marine fishes
Pisces
Predation
Predators
Sea water ecosystems
Synecology
title Predation, Prey Refuges, and the Structure of Coral-Reef Fish Assemblages
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