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Hay, Mark E.

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    Chemical defense in the seaweed Dictyopteris delicatula: differential effects against reef fishes and amphipods
    (Georgia Institute of Technology, 1988-09-21) Hay, Mark E. ; Duffy, J. Emmett ; Fenical, William ; Gustafson, Kirk
    Many seaweeds produce chemicals that deter feedlng by fishes and sea urchins. A growing body of evidence suggests that small, relatively immobile herbivores (mesograzers) such as amphpods, polychaetes, and ascoglossan gastropods are often unaffected by these compounds and may preferentially consume seaweeds that are chemically defended from fishes. We tested this hypothesis by examining the responses of reef fishes and amphipods to a mutture of 2 C,, hydrocarbons, &ctyopterenes A and B, produced by the Canbbean brown alga D~ctyopteris delicatula. This alga was intermediate in preference for reef fishes, and the dictyopterenes reduced fish grazing by a significant 40 %. In contrast, D. delicatula was highly preferred by a muted-species group of amphipods and the dlctyopterenes had no effect on their feeding Despite the tendency for mesograzers to selectively consume some seaweeds that are chemically deterrent to fishes, true specialization by these or other marine herbivores appears to be rare in companson with terrestnal systems. Plant-dwelling amphipods at our study site in the Grenadine Islands were found on, and consumed a variety of, macrophytes; they were not restrict~velys pecialized to D. delicatula. Many terrestnal insects are very specialized feeders, sequester toxins from theu food plants, and use these as duect defenses against predation. In contrast, sequestenng of seaweed toxlns by marine mesograzers appears to be relahvely rare. However, the indirect advantage of llving on seaweeds that are not eaten by fishes may be considerable. We hypothesize that mesograzers living on plants chemically defended from fishes wlll experience less predation than those living on plants preferred by fishes.
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    Chemical defense against different marine herbivores:  are amphipods insect equivalents? 
    (Georgia Institute of Technology, 1987) Hay, Mark E. ; Duffy, J. Emmett ; Pfister, Catherine A. ; Fenical, William
    The structurally similar diterpenoid alcohols pachydictyol-A and dictyol-E are produced by the brown seaweed Dictyota dichotoma. This seaweed and several related species that also produce these compounds are known to be relatively low preference foods for tropical fishes and urchins. We evaluated the effect of various concentrations of these compounds on feeding by the three common types of herbivores that co-occur with Dictyota in coastal North Carolina. Fish (Diplodus holbrooki), sea urchins (Arbacia punctulata), and a mixed species group of gammarid amphipods were offered pieces ofthe palatable seaweed Graci/aria tikvahiae coated with either (1) dictyol-E or pachydictyol-A dissolved in diethyl ether or (2) diethyl ether alone. Dictyol-E significantly reduced consumption by fish and urchins at concentrations of 0.5 and 1.0% of algal dry mass, but had no effect on amphipod grazing. Pachydictyol-A significantly reduced fish grazing at the relatively high concentrations of 1.0 and 1.3% of plant dry mass; at 0.5% it tended to decrease grazing, but the effect was not significant (P = .07). Pachydictyol-A had no effect on urchin grazing and significantly increased amphipod grazing. When Pachydictyol-A was fed to fish as 1.0% of food dry mass, their growth rate was reduced by a significant 48%. In feeding preference experiments with several seaweeds, Dictyota ranks low for fish and urchins but high for amphipods. This is consistent with the hypothesis that the secondary metabolites produced by Dictyota playa major role in determining its susceptibility to herbivores. The ability of amphipods to circumvent the chemical defenses of Dictyota, and the fact that the two species of algae most readily consumed by amphipods (Codium and Dictyota) were the two species least readily consumed by fish, suggest that predation and herbivory by fishes may be major factors selecting for amphipods that can live on, and eat, seaweeds that are unpalatable to fishes. Amphipods that fed on Dictyota did not appear to sequester the Dictyota metabolites; when exposed to fish predation, Dictyota-fed amphipods were eaten as readily as amphipods that had fed on an alga with no defensive chemistry. Tubicolous amphipods and other small marine herbivores that may spend significant portions of their lives on only a few plants may be under very different evolutionary constraints than the larger, more mobile herbivores that commonly move between many plants. Several characteristics of these smaller, less mobile, and much less studied, marine herbivores suggest that they may be ecologically similar to terrestrial insects and may play a large, but presently unappreciated, role in structuring marine plant communities.