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11th ICRS Abstract book - Nova Southeastern University

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Oral Mini-Symposium 10: Ecological Processes on Today's Reef Ecosystems<br />

10-37<br />

Fish Demography in The Large<br />

Richard VANCE* 1 , Mark STEELE 2 , Graham FORRESTER 3<br />

1 Ecology & Evolutionary Biology, UCLA, Los Angeles, CA, 2 Biology, California State<br />

<strong>University</strong> Northridge, Northridge, CA, 3 Natural Resources, <strong>University</strong> of Rhode Island,<br />

Kingston, RI<br />

Effective management of commercial coral reef fish metapopulations will require<br />

creation of marine protected areas. Informed design of MPA’s must employ<br />

demographic rate functions that drive fish metapopulation dynamics. Unfortunately,<br />

practical constraints usually force measurement of demographic rates on small portions<br />

of individual reefs, a spatial scale far too small to apply directly to metapopulations.<br />

Using these measurements therefore requires some way of “expanding” them to the<br />

spatial scale of entire large coral reefs.<br />

This talk will describe a practical scaling-up procedure. We will show, for the bridled<br />

goby Coryphopterus glaucofraenum (a convenient non-commercially fished species) how<br />

a whole-reef density-dependent death rate function can be constructed from an<br />

empirically measured death rate function that applies at the local scale. The procedure<br />

employs a computer simulation model that follows the fates of individual gobies in all 2m<br />

x 2m cells of a large reef. The reef habitat varies spatially both in its local density of<br />

crevices that provide refuge from predators to goby adults and in the substrate’s<br />

attractiveness to goby settlers. From simulated cell-level demographic data is calculated<br />

a whole-reef per capita death rate, and this rate is then statistically fitted to a function of<br />

whole-reef goby and refuge densities using maximum likelihood methods.<br />

Density dependence at the whole-reef scale is highly statistically significant and can be<br />

described by a function whose form resembles that of the local-scale function. However,<br />

reef spatial heterogeneity forces some adjustment in parameters. Indeed, density<br />

dependence at the whole-reef scale actually proves stronger than at the local scale. We<br />

shall display how scaling up the death rate function is influenced by variability in local<br />

refuge density and in local settler attraction, by the correlation of these reef attributes, and<br />

by the spatial scale on which this habitat heterogeneity occurs.<br />

10-38<br />

Large-Scale Experiment Reveals Effects Of Habitat Structure On Coral Reef Fish<br />

Assemblages<br />

Mark STEELE* 1 , Graham FORRESTER 2 , Jameal SAMHOURI 3 , Clare WORMALD 2<br />

1 Department of Biology, California State <strong>University</strong> Northridge, Northridge, CA,<br />

2 Department of Natural Resources Science, <strong>University</strong> of Rhode Island, Kingston, RI,<br />

3 Northwest Fisheries Science Center, NOAA Fisheries, Seattle, WA<br />

The structure of coral reef fish assemblages is usually strongly correlated with attributes<br />

of reef structure. Experimental studies on small patch reefs have improved our<br />

understanding of how certain attributes of reef structure (e.g., shelter hole size) influence<br />

these assemblages, but it is not clear if these results from small-scale experiments<br />

extrapolate to larger spatial and temporal scales. We manipulated the abundance of<br />

rubble habitat to evaluate how this shelter-providing habitat affected the structure of coral<br />

reef fish assemblages at large scales. We added several tons of limestone rubble and<br />

conch shells to one half of each of 5 large (3,000-15,000 m 2 ), isolated reefs near Lee<br />

Stocking Island, Bahamas. We tracked changes in the reef fish assemblage with<br />

underwater visual censuses over 4 years. On rubble addition halves of reefs, several<br />

species of small fishes (damselfishes, wrasses, and gobies) became more abundant,<br />

whereas several similar species in the same families were unaffected. Small-scale habitat<br />

associations and results of previous small-scale habitat manipulations generally did a<br />

good job of predicting which species would be affected by the habitat manipulation and<br />

which were not. Larger species (e.g., snappers, grouper, and squirrelfishes) were<br />

unaffected by the manipulation, presumably because they are not dependent on small<br />

crevices for shelter and their abundance is not influenced by the abundance of smaller<br />

potential prey species that increased as a result of the rubble addition. There was no<br />

strong evidence that any species was negatively affected by the habitat manipulation.<br />

Thus, our results indicate that assemblage structure can by altered by large-scale habitat<br />

manipulations, which increase the abundance of some habitat-dependent species, without<br />

unintended negative effects on other species.<br />

10-39<br />

Extrapolating From Small-Scale Experiments: Predation And Refuge-Shortage in Coral<br />

Reef Fishes<br />

Graham FORRESTER* 1 , Mark STEELE 2 , Jameal SAMHOURI 3 , Richard VANCE 4<br />

1 <strong>University</strong> of Rhode Island, Kingston, RI, 2 Biology, California State <strong>University</strong> Northridge,<br />

Northridge, CA, 3 NOAA Fisheries, Seattle, WA, 4 <strong>University</strong> of California Los Angeles, Los<br />

Angeles, CA<br />

Although field experiments allow rigorous tests of ecological hypotheses, they are usually<br />

limited to small spatial scales. We often want to know if their findings extrapolate to larger<br />

scales, especially when seeking to apply their results to conservation and management.<br />

Experiments on small coral reef fishes (bridled gobies) occupying small habitat patches reveal<br />

that locally density-dependent mortality is inflicted by predators. As prey become crowded,<br />

they suffer a progressively increasing shortage of structural refuges. Consequently, goby<br />

mortality at small-scales is well described by a model in which vulnerability is proportional to<br />

the ratio of gobies to refuges. A manipulation of refuge abundance on entire reefs, which are<br />

the size of small marine reserves and approach the scale at which some reef fisheries operate,<br />

suggests that a similar interaction occurs at this much larger spatial scale. This result is in<br />

accord with a scaling model which indicates that the effects of refuge shortage on entire reefs<br />

can be extrapolated from the model describing goby vulnerability as proportional to the ratio of<br />

fish to refuges. In simple terms, the biological interactions causing density dependence scale<br />

up. Careful extrapolation from small-scale experiments identifying species-interactions may<br />

thus be possible, and so should improve our ability to predict the outcomes of alternate<br />

management strategies for coral reef fishes.<br />

10-40<br />

Musical chairs: competition for unguarded refuges and density-dependent mortality<br />

Jameal SAMHOURI* 1,2 , Richard VANCE 3 , Graham FORRESTER 4 , Mark STEELE 5<br />

1 Northwest Fisheries Science Center, NOAA Fisheries, Seattle, WA, 2 Department of Ecology<br />

and Evolutionary Biology, UC Los Angeles, Los Angeles, 3 Department of Ecology and<br />

Evolutionary Biology, UC Los Angeles, Los Angeles, CA, 4 Department of Natural Resources<br />

Science, <strong>University</strong> of Rhode Island, Kiingston, RI, 5 Department of Biology, California State<br />

<strong>University</strong> Northridge, Northridge, CA<br />

Predation is widely recognized as a dominant structuring force in marine populations and<br />

communities. As a result, refuges that reduce predation risk can be an important limiting<br />

resource in prey populations, ultimately mediating the occurrence and strength of densitydependent<br />

mortality. On coral reefs, many fishes retreat to unguarded structural refuges, such as<br />

crevices under coral or other hard substrate, only in response to a predator strike. This behavior<br />

leads to a type of competition for refuges that resembles conceptually the childhood game of<br />

musical chairs: prey scramble to find a safe place to hide when threatened or attacked, and only<br />

when refuges are abundant relative to the number of retreating prey is it likely that individuals<br />

will successfully evade predation. We introduce a new mathematical function to describe the<br />

relationship between structural refuges, population density, and per capita mortality in bridled<br />

gobies (Coryphopterus glaucofraenum) under this form of competition. In this model, we<br />

assume that the proximate cause of all goby deaths is predation, a fixed fraction of individuals<br />

do not respond to predator strikes by retreating to structural refuges, and the remaining<br />

individuals are capable of seeking shelter in a fixed number of unguarded refuges when<br />

threatened or attacked. Therefore, the likelihood of an individual dying during a predation event<br />

is either constant and density-independent or directly proportional to refuge availability, and<br />

these two components of mortality are not additive. In comparison to conventional densitydependent<br />

mortality functions, our musical chairs function provides a much improved fit to<br />

available empirical data for bridled gobies. We suggest that the musical chairs function may<br />

have broad applicability to other coral reef fishes, and more generally, for species in which<br />

refuges from predation are a limiting resource but territorial refuge-guarding does not occur.<br />

82

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