Does the skeleton of a sponge provide a defense against predatory reef fish?

Sponge tissue often contains two structural components in high concentrations: spicules of silica, and refractory fibers of protein (spongin). Some terrestrial plants contain analogous structures, siliceous inclusions and refractory lignins, that have been demonstrated to deter herbivory. We performed feeding experiments with predatory reef fish to assess the deterrent properties of the structural components of three common Caribbean demosponges, Agelas clathrodes, Ectyoplasia ferox, and Xestospongia muta. The concentrations of spicules and spongin in the tissues varied widely between the three species, but when assayed at their natural volumetric concentrations, neither spicules (all three species assayed) nor the intact spiculated spongin skeleton (A. clathrodes and X. muta assayed) deterred feeding by reef fish in aquarium or field assays using prepared foods of a nutritional quality similar to, or higher than, that of sponge tissue. Spicules deterred feeding in aquarium assays when incorporated into prepared foods of a nutritional quality lower than that of sponge tissue (15–19 times less protein), but spiculated spongin skeleton was still palatable, even in prepared foods devoid of measurable protein, and even though spicules embedded in spongin were oriented in their natural conformation. Based on comparisons of the nutritional qualities of the tissues of the three sponge species and of the prepared foods, sponge tissue would have to be much lower in food value (5 times less protein or lower) for spicules to provide an effective defense, and spicules in combination with the spongin skeleton would be unlikely to provide an effective defense regardless of the nutritional quality of the tissue. Unlike terrestrial plants, marine sponges may use silica and refractory fibers solely for structural purposes.     

Within-colony feeding selectivity by a corallivorous reef fish: foraging to maximize reward?

Foraging theory predicts that individuals should choose a prey that maximizes energy rewards relative to the energy expended to access, capture, and consume the prey. However, the relative roles of differences in the nutritive value of foods and costs associated with differences in prey accessibility are not always clear. Coral‐feeding fishes are known to be highly selective feeders on particular coral genera or species and even different parts of individual coral colonies. The absence of strong correlations between the nutritional value of corals and prey preferences suggests other factors such as polyp accessibility may be important. Here, we investigated within‐colony feeding selectivity by the corallivorous filefish, Oxymonacanthus longirostris, and if prey accessibility determines foraging patterns. After confirming that this fish primarily feeds on coral polyps, we examined whether fish show a preference for different parts of a common branching coral, Acropora nobilis, both in the field and in the laboratory experiments with simulated corals. We then experimentally tested whether nonuniform patterns of feeding on preferred coral species reflect structural differences between polyps. We found that O. longirostris exhibits nonuniform patterns of foraging in the field, selectively feeding midway along branches. On simulated corals, fish replicated this pattern when food accessibility was equal along the branch. However, when food access varied, fish consistently modified their foraging behavior, preferring to feed where food was most accessible. When foraging patterns were compared with coral morphology, fish preferred larger polyps and less skeletal protection. Our results highlight that patterns of interspecific and intraspecific selectivity can reflect coral morphology, with fish preferring corals or parts of coral colonies with structural characteristics that increase prey accessibility.     

Energy flow to two abundant consumers in a subtropical oyster reef food web

Oyster reefs are among the most threatened coastal habitat types, but still provide critical habitat and food resources for many estuarine species. The structure of oyster reef food webs is an important framework from which to examine the role of these reefs in supporting high densities of associated fishes. We identified major trophic pathways to two abundant consumers, gray snapper (Lutjanus griseus) and crested goby (Lophogobius cyprinoides), from a subtropical oyster reef using stomach content and stable isotope analysis. The diet of gray snapper was dominated by crabs, with shrimp and fishes also important. Juvenile gray snapper fed almost entirely on oyster reef-associated prey items, while subadults fed on both oyster reef- and mangrove-associated prey. Based on trophic guilds of the gray snapper prey, as well as relative δ¹³C values, microphytobenthos is the most likely basal resource pool supporting gray snapper production on oyster reefs. Crested goby had omnivorous diets dominated by bivalves, small crabs, detritus, and algae, and thus were able to take advantage of prey relying on production from sestonic, as well as microphytobenthos, source pools. In this way, crested goby represent a critical link of sestonic production to higher trophic levels. These results highlight major trophic pathways supporting secondary production in oyster reef habitat, thereby elucidating the feeding relationships that render oyster reef critical habitat for many ecologically and economically important fish species.     

Seaweed-herbivore-predator interactions: host-plant specialization reduces predation on small herbivores

Summary Because feeding specialization among marine herbivores is rare, marine communities provide a simplified system for identifying factors selecting for specialization. On Australia's Great Barrier Reef, we investigated interactions among the chemically-defended seaweed Chlorodesmis fastigiata, herbivores specialized on this alga, and potential predators of these herbivores. Chlorodesmis is a low preference food for reef fishes but appears to be the only food of the crab Caphyra rotundifrons and the ascoglossan gastropods Elysia sp. and Cyerce nigricans. The crab is found only in patches of Chlorodesmis, feeds solely on the alga, and selectively shelters in it in laboratory choice experiments. Crab grazing on the red seaweed Acanthophora spicifera was stimulated when this alga was coated with increasing concentrations of the cytotoxic diterpenoid chlorodesmin, the major secondary metabolite of Chlorodesmis. Crabs did not sequester Chlorodesmis metabolites but avoided predators by sheltering in the unpalatable alga. All crabs tethered on the reef without access to Chlorodesmis patches were rapidly eaten; those with access to Chlorodesmis patches were much less susceptible to predation. The cryptic ascoglossan Elysia sp. was found exclusively in patches of Chlorodesmis and sequestered metabolites from the alga. Living Elysia were unpalatable to the common wrasse Thalassoma lunare in laboratory assays, but the crude organic extract of Elysia did not significantly deter feeding by Thalassoma. Elysia sequestered chlorodesmin, which deterred feeding by reef fishes in field assays but was ineffective against Thalassoma in laboratory assays at 5% food dry mass. Unlike Elysia, the aposematically colored ascoglossan Cyerce nigricans sequestered Chlorodesmis metabolites in relatively small amounts, but produced larger amounts of unrelated polypropionate compounds. Cyerce were never attacked by fishes and the crude organic extract of this slug strongly deterred feeding by wrasses in laboratory assays. The dorid nudibranch Gymnodoris sp. was found only in Chlorodesmis patches and appeared to be a specialized predator on Elysia; it would not prey on Cyerce. Data from this and other recent investigations demonstrate that some small marine herbivores feed selectively or exclusively on seaweeds that are chemically defended from fishes. This association reduces predation on the herbivores and suggests that escape from and deterrence of predation may be a dominant factor selecting for specialization among these herbivores.     

Chemical and structural defenses in the sea fan Gorgonia ventalina: effects against generalist and specialist predators

Cyphoma gibbosum is an ovulid predatory gastropod that specializes on gorgonians, many of which contain secondary metabolites and calcified sclerites that serve as antipredator defenses. In field and shipboard feeding assays, we examined the role of gorgonian crude extracts and sclerites as feeding deterrents to generalist predators and to C. gibbosum. Crude extracts and sclerites were isolated from Gorgonia ventalina, a Caribbean sea fan on which C. gibbosum feeds, and incorporated into a carrageenan-based artificial diet. In shipboard feeding experiments, artificial diet containing G. ventalina crude extracts was consumed 49% less by C. gibbosum, than artificial diet lacking extracts. The addition of G. ventalina sclerites to the diet also reduced feeding by Cyphoma by about one half. The addition of extracts to the artificial diet reduced feeding by natural assemblages of tropical fishes at Cross Harbor, Great Abaco Island, Bahamas by 87%; sclerites reduced feeding by fishes by 95%. Gorgonia ventalina extracts were composed of at least a dozen nonpolar terpenoids. Fractions containing these compounds were feeding deterrents towards fishes in the field. Unlike many terrestrial oligophagous specialists, C. gibbosum is not immune to the defenses produced by its prey.     

Large mobile versus small sedentary herbivores and their resistance to seaweed chemical defenses

Summary Small, relatively sedentary herbivores like amphipods and polychaetes (mesograzers) often live on the plants they consume and should therefore view plants as both foods and living sites. Large, relatively mobile herbivores like fishes commonly move among, and feed from, many plants; they should view plants primarily as foods and rarely as potential living sites. In marine communities, fishes that consume plants are also important predators on mesograzers. Since seaweeds avoided by fishes should represent safer living sites for small herbivores, mesograzers living on and consuming seaweeds that are not eaten by fishes should have higher fitness than mesograzers living on plants preferred by fishes. In previous work, we demonstrated that seaweed secondary metabolites that deterred feeding by a fish and sea urchin had no effect on feeding by a common amphipod (Hay et al. 1987a). We then hypothesized that mesograzers would, in general, be less affected by seaweed chemical defenses than larger, more mobile herbivores like fishes. In this investigation, we evaluate the generality of this hypothesis by comparing the feeding of an omnivorous fish (Lagodon rhomboides) with that of an omnivorous, tube-building polychaete (Platynereis dumerilii) to see if the mesograzer prefers seaweeds avoided by the fish and if it is less affected by seaweed chemical defense. Platynereis dumerilii fed almost exclusively on Dictyota dichotoma, the seaweed eaten least by Lagodon rhomboides. The diterpene alcohols (dictyol-E and pachydictyol-A) produced by Dictyota significantly deterred feeding by Lagodon but did not affect, or at one concentration stimulated, feeding by Platynereis. Our data support the hypothesis that small, relatively sedentary herbivores that live on plants are more resistant to chemical defenses than are large, relatively mobile herbivores that move among many plants.     

Sponge community structure and anti-predator defenses on temperate reefs of the South Atlantic Bight

Predator–prey interactions can play a significant role in shaping the structure of both terrestrial and marine communities. Sponges are major contributors to benthic community structure on temperate reefs and although several studies have investigated how abiotic processes control sponge distributions on these reefs, the role of predation is less clear. We investigated the relationship between sponge predators and the distribution of sponges on temperate reefs in the South Atlantic Bight (SAB), off Georgia, USA. We documented sponge species richness and abundance, spongivorous fish density, and examined the ability of 19 sponge species to chemically and structurally deter predation by fishes. We also conducted reciprocal transplant experiments to determine if predation by fishes contributes to the observed zonation of sponge species on these reefs. Our surveys revealed two distinct sponge assemblages: one characterized by amorphous and encrusting sponge morphotypes colonizing the vertical, rocky outcroppings (scarp sponge community), while the other consisted of pedunculate, digitate, and arborescent growth forms occurring on the sediment-laden reef top (plateau sponge community). Spongivorous fishes were more abundant on the scarp than the plateau and scarp sponges were found to be more effective than plateau sponges at chemically deterring generalist fishes. In contrast, plateau sponges were more reliant on structural defenses: a result consistent with the higher spicule content of their skeletons. Transplant experiments confirmed that predators prevent some plateau sponges from colonizing the scarp even though they possess structural defenses. Thus, predation appears to play a role in shaping sponge community structure on SAB reefs by restricting those species lacking adequate chemical defenses to habitats where there is a paucity of spongivores.     

Effects of seaweed extracts and secondary metabolites on feeding by the herbivorous surgeonfish Naso lituratus

We examined 22 species of algae and two species of seagrasses from coral reef habitats around Guam to determine if they possessed chemical defenses against the acanthurid Naso lituratus. Whole plants (18 species) were offered to determine whether they were preferred or avoided by N. lituratus in the laboratory. Organic extracts of 15 algae and one seagrass were applied to palatable seaweeds and offered to N. lituratus in the laboratory to determine if the seaweeds were chemically defended. Extracts that deterred feeding were further fractionated if sufficient amounts were available, and the fractions and associated pure compounds were tested in similar feeding assays. N. lituratus was significantly deterred from feeding by crude extracts from five different species of algae: Avrainvillea obscura, Bryopsis pennata, grazed Halimeda macroloba, Neomeris annulata, and Portieria (=Desmia) hornemannii. The pure compounds avrainvilleol from A. obseura, ochtodene from P. hornemannii, one fraction and one brominated sesquiterpene from N. annulata, and two fractions from T. expeditionis also deterred feeding. These results, together with previous work, suggest that tropical herbivorous fishes differ in their responses to plant chemistry, and this variability precludes broad generalization about the effects of marine plant secondary metabolites on herbivorous fishes.     

Testing for defensive synergy in Caribbean sponges: Bad taste or glass spicules?

Chemical and physical defenses of sessile organisms against consumers are well described for both terrestrial and marine systems. However, previous studies have focused on chemical or physical defenses in isolation, and have not considered their interaction. Marine sponges provide a model system for testing this interaction. Some sponge species produce secondary metabolites that deter predation; they may also contain siliceous spicules, but previous studies have provided little evidence that spicules in isolation offer any defense against generalist fish predators. To determine whether the two components have an additive, antagonistic, or synergistic interaction, crude organic extracts and spicules from individuals of 8 Caribbean sponge species were isolated and tested in laboratory feeding assays. These included one chemically defended reef sponge (Agelas clathrodes) and seven known to be intermediately deterrent: six from reef habitats (Cinachyrella alloclada, Clathria virgultosa, Cribrochalina infundibulum, Niphates digitalis, Svenzea zeai, and Xestospongia muta) and one from mangrove habitats (Tedania ignis). Extracts and spicules were assayed at various concentrations, both individually and in combination, in laboratory feeding assays with the bluehead wrasse, Thalassoma bifasciatum. A SAS based GENMOD procedure based on an isobolographic analysis model was used for statistical comparisons. Four sponges (A. clathrodes, C. alloclada, C. virgultosa, and one of three individuals of X. muta) showed evidence of synergisms. Of these, synergy in C. alloclada, C. virgultosa, and X. muta was caused by approximately natural concentrations of extracts and spicules. The extract of A. clathrodes was deterrent, but combination assays required nearly a 3-fold reduction in extract concentration and an 8-fold increase in spicule concentration to show the synergistic effect. Contrary to previous findings, spicules from C. infundibulum and two of three individuals of X. muta were deterrent at natural concentrations. Sponge spicules may be defensive in isolation, or may enhance chemical defenses against consumers, but the lack of synergisms for individuals in 4 of 7 species with intermediate levels of chemical defense suggests that defensive synergy is not the general rule and, when present, may be an example of an exaptation.     

Food partitioning and spatial subsidy in shelter‐limited fishes inhabiting patchy reefs of Patagonia

The diets of the most conspicuous reef‐fish species from northern Patagonia, the carnivorous species Pseudopercis semifasciata, Acanthistius patachonicus, Pinguipes brasilianus and Sebastes oculatus were studied. Pinguipes brasilianus had the narrowest diet and most specialized feeding strategy, preying mostly on reef‐dwelling organisms such as sea urchins, limpets, bivalves, crabs and polychaetes. The diet of A. patachonicus was characterized by the presence of reef and soft‐bottom benthic organisms, mainly polychaetes, crabs and fishes. Pseudopercis semifasciata showed the broadest spectrum of prey items, preying upon reef, soft‐bottom and transient organism (mainly fishes, cephalopods and crabs). All S. oculatus guts were empty, but stable‐isotope analyses suggested that this species consumed small fishes and crabs. In general, P. brasilianus depended on local prey populations and ate different reef‐dwelling prey than the other species. Pseudopercis semifasciata, A. patachonicus and probably S. oculatus, however, had overlapping trophic niches and consumed resources from adjacent environments. The latter probably reduces the importance of food as a limiting resource for these reef‐fish populations, facilitating their coexistence in spite of their high trophic overlap.     

Food selection by calanoid copepods in response to between-lake variation in food abundance

1. Food selection experiments were conducted by acclimating calanoid copepods (Eudiaptomus spp.) in suspensions of natural seston and then adding pairs of dual-labelled (¹⁴C/³²P) algae. Each feeding trial measured selectivity between a small, high-quality reference alga, Chlamydomonas reinhardii, and a test alga that differed in size and/or food quality. The influence of food concentration on food selection was tested by using seston from two lakes with contrasting food abundance and by including treatments with filtered lake water ('starved’) and seston diluted with filtered water or enriched with cultured algae. 2. Copepods that had been starved or acclimated to natural seston with low food abundance preferred the larger of two labelled algae, regardless of the nutritional quality of the algae. In agreement with the predictions of an optimal diet model, however, copepods that were acclimated to high food conditions discriminated against low-quality foods, including digestion-resistant algae and dead algae. 3. Selectivity coefficients showed excellent agreement with a previous study involving the same taxa of copepods and labelled algae but in which the copepods had been acclimated to pairs of cultured algae rather than natural seston. Thus, these comparisons emphasize the importance of food availability in modulating copepod selectivity for foods that differ in nutritional quality and suggest that such behaviour occurs in nature.     

The importance of honeydew as food for larvae of Chrysoperla carnea in the presence of aphids

Larvae of the common green lacewing Chrysoperla carnea are predacious and feed on a wide range of small, soft‐bodied arthropods. In addition to their feeding on prey arthropods to cover their nutritional requirements for growth and development, the consumption of non‐prey foods such as honeydew has been reported. It is commonly believed that these food supplements are primarily exploited by the larvae when prey is scarce or of low nutritional quality. Here, we assess whether C. carnea larvae also use honeydew when high‐quality aphid prey are readily available. In a choice experiment, the feeding behaviour of C. carnea larvae was observed in the presence of both aphids and honeydew. The larvae were starved, aphid‐fed, or honeydew‐fed prior to the experiment. The time spent feeding on honeydew compared with feeding on aphids was highest for starved larvae and lowest for honeydew‐fed larvae. Among the three treatments, the aphid‐fed larvae spent the most time resting and the least time searching. In an additional experiment food intake was assessed in terms of weight change when larvae were provided with an ad libitum supply of either aphids or honeydew. Larvae yielded a significant lower relative weight increase on honeydew compared with aphids. The reduced weight increase on honeydew was compensated when larvae were subsequently provided with aphids, but not when honeydew was provided again. This study showed that (i) prior honeydew feeding reduces overall aphid consumption, and (ii) larvae do consume honeydew even after they have been given ad libitum access to aphids. The fact that larvae of C. carnea still use honeydew as a food source in the presence of suitable prey underlines the importance of carbohydrates as foods.     

Intentional partial beaching in a coral reef fish: a newly recorded hunting behaviour of titan triggerfish,Balistoides viridescens

Coral reef fishes use a multitude of diverse feeding behaviours to increase their ability to successfully capture a wide range of prey. Here, this study reports a novel hunting behaviour in a coral reef fish, the titan triggerfish, Balistoides viridescens, where an individual was seen partially beaching itself while attempting to catch a Red Sea ghost crab, Ocypode saratan. This is the first report of this behaviour in the order Tetraodontiformes and represents an astonishing capability of this species to exploit food resources outside their typical assumed ecological niche.     

Trophic relationships among concentrated populations of small fishes in seagrass meadows

Changes in available food and utilized foods for a densely populated guild of animals can uncover periods of niche displacement among particular consumer species or their separate size classes. Dense populations of small fishes and their foods were çensussed for 13 months and studied experimentally in shoalgrass and turtlegrass meadows of Redfish Bay, Texas. Feeding habits were determined with respect to prey availability, and illustrated the extent to which seasonal partitioning of food corresponded to food depletion among these abundant consumers in seagrass meadow food webs. The darter goby, code goby, pinfish, and Gulf pipefish were the most common species throughout the year. Although the darter goby did not show distinct ontogenetic changes in food habits, the code goby, Gulf pipefish, and pinfish demonstrated major ontogenetic progressions of foods selected.Food availability in the seagrass meadows changed seasonally. When major prey such as amphipods were abundant, during spring, many fish species showed high overlap in food use. Regardless of food availability, the code goby and Gulf pipefish fed mainly on amphipods or copepods. The more common darter goby and pinfish seasonally changed their diets with changes in food availability; the darter goby and pinfish were more carnivorous during spring, but they largely consumed epiphytic algae during summer. Cage experiments were used to monitor foods confined with elevated densities of pinfish and darter gobies, relative to control cages at natural overall densities. Prey items in the former cages decreased sharply, with corresponding dietary shifts by these common fishes. During resource depletion, these changes in resource use by these naturally concentrated consumers appeared as temporary partitioning of available resources. This shift occurred during both natural (seasonal) and experimental depletions of food, and appears to result from increased interspecific and intraspecific competition during periods of depleted preferred foods.     

Simulated eutrophication and browning alters zooplankton nutritional quality and determines juvenile fish growth and survival

The first few months of life is the most vulnerable period for fish and their optimal hatching time with zooplankton prey is favored by natural selection. Traditionally, however, prey abundance (i.e., zooplankton density) has been considered important, whereas prey nutritional composition has been largely neglected in natural settings. High‐quality zooplankton, rich in both essential amino acids (EAAs) and fatty acids (FAs), are required as starting prey to initiate development and fast juvenile growth. Prey quality is dependent on environmental conditions, and, for example, eutrophication and browning are two major factors defining primary producer community structures that will directly determine the nutritional quality of the basal food sources (algae, bacteria, terrestrial matter) for zooplankton. We experimentally tested how eutrophication and browning affect the growth and survival of juvenile rainbow trout (Oncorhynchus mykiss) by changing the quality of basal resources. We fed the fish on herbivorous zooplankton (Daphnia) grown with foods of different nutritional quality (algae, bacteria, terrestrial matter), and used GC‐MS, stable isotope labeling as well as bulk and compound‐specific stable isotope analyses for detecting the effects of different diets on the nutritional status of fish. The content of EAAs and omega‐3 (ω‐3) polyunsaturated FAs (PUFAs) in basal foods and zooplankton decreased in both eutrophication and browning treatments. The decrease in ω‐3 PUFA and especially docosahexaenoic acid (DHA) was reflected to fish juveniles, but they were able to compensate for low availability of EAAs in their food. Therefore, the reduced growth and survival of the juvenile fish was linked to the low availability of DHA. Fish showed very low ability to convert alpha‐linolenic acid (ALA) to DHA. We conclude that eutrophication and browning decrease the availability of the originally phytoplankton‐derived DHA for zooplankton and juvenile fish, suggesting bottom‐up regulation of food web quality.     

Novel species interactions and environmental conditions reduce foraging competency at the temperate range edge of a range-extending coral reef fish

Poleward range extensions of coral reef species can reshuffle temperate communities by generating competitive interactions that did not exist previously. However, novel environmental conditions and locally adapted native temperate species may slow tropical invasions by reducing the ability of invaders to access local resources (e.g. food and shelter). We test this hypothesis on wild marine fish in a climate warming hotspot using a field experiment encompassing artificial prey release. We evaluated seven behaviours associated with foraging and aggressive interactions in a common range-extending coral reef fish (Abudefduf vaigiensis) and a co-shoaling temperate fish (Microcanthus strigatus) along a latitudinal temperature gradient (730 km) in SE Australia. We found that the coral reef fish had reduced foraging performance (i.e. slower prey perception, slower prey inspection, decreased prey intake, increased distance to prey) in their novel temperate range than in their subtropical range. Furthermore, higher abundance of temperate fishes was associated with increased retreat behaviour by coral reef fish (i.e. withdrawal from foraging on released prey), independent of latitude. Where their ranges overlapped, temperate fish showed higher foraging and aggression than coral reef fish. Our findings suggest that lower foraging performance of tropical fish at their leading range edge is driven by the combined effect of environmental factors (e.g. lower seawater temperature and/or unfamiliarity with novel conditions in their extended temperate ranges) and biological factors (e.g. increased abundance and larger body sizes of local temperate fishes). Whilst a future increase in ocean warming is expected to alleviate current foraging limitations in coral reef fishes at leading range edges, under current warming native temperate fishes at their trailing edges appear able to slow the range extension of coral reef fishes into temperate ecosystems by limiting their access to resources.

     

Chemical defense of the eastern newt (Notophthalmus viridescens): variation in efficiency against different consumers and in different habitats

Amphibian secondary metabolites are well known chemically, but their ecological functions are poorly understood--even for well-studied species. For example, the eastern newt (Notophthalmus viridescens) is a well known secretor of tetrodotoxin (TTX), with this compound hypothesized to facilitate this salamander's coexistence with a variety of aquatic consumers across the eastern United States. However, this assumption of chemical defense is primarily based on observational data with low replication against only a few predator types. Therefore, we tested the hypothesis that N. viridescens is chemically defended against co-occurring fishes, invertebrates, and amphibian generalist predators and that this defense confers high survivorship when newts are transplanted into both fish-containing and fishless habitats. We found that adult eastern newts were unpalatable to predatory fishes (Micropterus salmoides, Lepomis macrochirus) and a crayfish (Procambarus clarkii), but were readily consumed by bullfrogs (Lithobates catesbeianus). The eggs and neonate larvae were also unpalatable to fish (L. macrochirus). Bioassay-guided fractionation confirmed that deterrence is chemical and that ecologically relevant concentrations of TTX would deter feeding. Despite predatory fishes rejecting eastern newts in laboratory assays, field experiments demonstrated that tethered newts suffered high rates of predation in fish-containing ponds. We suggest that this may be due to predation by amphibians (frogs) and reptiles (turtles) that co-occur with fishes rather than from fishes directly. Fishes suppress invertebrate consumers that prey on bullfrog larvae, leading to higher bullfrog densities in fish containing ponds and thus considerable consumption of newts due to bullfrog tolerance of newt chemical defenses. Amphibian chemical defenses, and consumer responses to them, may be more complex and indirect than previously appreciated.     

Influence of Trophic Relations on Form and Behavior Among Fishes and Benthic Invertebrates in Some California Marine Communities

Study of interactions among fishes and benthic invertebrates off southern California's Santa Catalina Island showed the great extent to which taxa are defined by trophic relations. The dominant fishes there are acanthopterygian teleosts, with the serranine serranid Paralabrax clathratus morphologically nearest the evolutionary mainstream. Because mainstream species have morphologies similar to their progenitors, marine communities have a long history of adaptations to threats from generalized predators like P. clathratus. Species examined at Santa Catalina indicate this experience has produced four basic defenses: smallness, which presents difficulties for predators with large and simply constructed feeding mechanisms; dissemblance, as through camouflage, which makes detection by visual hunters difficult; inedibility, which is attained by incorporating materials that an unspecialized digestive system cannot accommodate; and nocturnality, which results in avoiding diurnal predators altogether. Examination of trophic relations among the major fishes in Santa Catalina communities shows each with specialized ways to counter these defenses. Prey smallness has been met by reducing size of mouth relative to size of prey, either by evolving as smaller fishes or through structural changes in head and jaws. Prey dissemblance has been accommodated through enhanced visual acuity and varied means to remain an unrecognized threat while waiting for prey to move. Inedibility has been countered by developing specialized features of the alimentary tract, including dentition, while adjustments to nocturnality have included enhanced sensory capabilities that detect prey in low light. These adaptations to prey defenses define varied lines of divergence from the teleost mainstream as represented atbreak Santa Catalina.     

Plant and Prey Consumption Cause a Similar Reductions in Cannibalism by an Omnivorous Bug

Cannibalism is usually more intense when other food sources are scarce, be it prey or plant-based foods. We hypothesized that feeding on plants would reduce cannibalism to a lesser extent than feeding on prey, because plants are considered nutritionally inferior compared to prey. We used the omnivorous bug Orius laevigatus Say (Heteroptera: Anthocoridae) to test this prediction. Starved female bugs were individually held with five second-instar conspecific nymphs and offered (i) Helicoverpa armigera eggs (prey); (ii) pollen (plant); (iii) H. armigera eggs and pollen (prey+plant); or (iv) no eggs or pollen. Fewer cannibalistic events and shorter feedings on conspecifics were recorded in the presence of pollen, prey or both than in their absence. Data therefore do not support our hypothesis that cannibalism is differentially affected by foods of different nutritional values. It seems that omnivorous feeding habits enable predators to sustain themselves on plant sources in the absence of prey, without the need to resort to cannibalism.     

Hydroid defenses against predators: the importance of secondary metabolites versus nematocysts

Marine hydroids are commonly thought to be defended by stinging organelles called nematocysts that penetrate predator tissues and inject proteinaceous venoms, but not all hydroids possess these nematocysts. Although an increasing number of bioactive secondary metabolites have been isolated from marine hydroids, ecological roles of these compounds are poorly known. To test the hypothesis that nematocysts and noxious secondary metabolites represent alternative defenses against predation, we examined hydroids from North Carolina, United States for: (1) the palatability of whole polyps before and after nematocysts had been deactivated; (2) the palatability of their chemical extracts; and (3) their nutritional value in terms of organic content, protein content, and levels of refractory structural material (chitin). All hydroids were avoided by a generalist predator, the pinfish Lagodon rhomboides, compared with palatable control foods. Two of these (Halocordyle disticha and Tubularia crocea) became palatable after being treated with potassium chloride to discharge their nematocysts, suggesting that these species rely on nematocysts for defenses against predators. Chemical extracts from nematocyst-defended species had no effect on fish feeding. The four species that remained unpalatable after nematocysts had been discharged (Corydendrium parasiticum, Eudendrium carneum, Hydractinia symbiolongicarpus, Tridentata marginata) possessed chemical extracts that deterred feeding by pinfish. We have isolated and characterized the structures of the deterrent metabolites in two of these species. We found no differences in nutritional content or levels of chitin between nematocyst-defended and chemically defended species, and no evidence that either of these played a role in the rejection of hydroids as prey. Our results suggest that, among hydroids, chemical defenses may be at least as common as nematocyst-based defenses and that the two may represent largely alternative defensive strategies. The four hydroid species with deterrent extracts represent four families and both sub-orders of hydroids, suggesting that chemical defenses in this group may be widespread and have multiple origins. Received: 25 May 1999 / Accepted: 1 February 2000