Published research about wetland insects has proliferated, and a conceptual foundation about how wetland insect populations and communities are regulated is being built. Here we review and synthesize this new body of work. Our review begins with a summary of insect communities found in diverse wetland types, marshes, forested floodplains, and peatlands. Next, we critically discuss research on the population and community ecology of wetland insects, including the importance of colonization strategies and insect interactions with the physical environment, plants, predators, and competitors. Results from many of the experimental studies that we review indicate that some commonly held beliefs about wetland insect ecology require significant reevaluation. We then discuss the importance of wetland insect ecology for some applied concerns such as efforts to manage wetland insect resources as waterfowl food and development of ecologically sound strategies to control pest mosquitoes. We conclude with a discussion of wetland conservation, emphasizing insect aspects.

We conducted manipulative field experiments in artificial ponds to quantify the predatory impact of larvae of a migratory dragonfly (Tramea lacerata) on a common resident dragonfly species (Erythemis simplicicollis), and on damselflies as shared prey of the two dragonflies. We found that the combined predatory effects of these two dragonflies on damselflies were not additive. To determine the underlying cause of non—additive predation rates in the field, we conducted a second experiment in laboratory aquaria to isolate the impact of each predator on the consumption rates of the other. Dragonfly consumption rates of damselflies in single—predator treatments were compared to those in the presence of heterospecifics or conspecifics with their menta (mouthparts) surgically modified so that they could not capture prey. In the laboratory experiment, de—mented Tramea reduced the consumption rates of Erythemis to less than half of that observed when Erythemis foraged alone. Erythemis numbers were also reduced by Tramea predation. Erythemis had neither effect on Tramea. Both of the negative effects of Tramea on Erythemis will have indirect positive effects on damselflies. The "behavioral" component (reduced Erythemis foraging rate) should be more important than the "trophic link" (reduced Erythemis numbers) indirect effect. Together these indirect positive effects will allay, but not completely compensate for, the direct negative effects of Tramea predation on damselflies. These results illustrate how an asymmetric potential for intraguild predation can lead to asymmetries in interference competition and to non—additive effects on prey mortality. The addition of removal of predators that interact in this manner to or from communities should have only a small net effect on prey because of compensating direct and indirect effects. This may explain why predator manipulations have often had unpredictable or undetectable effects on freshwater benthic communities.

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SUMMARY 1. Larvae of cased caddisflies (Limnephilidae and Phryganeidae) are among the most abundant and conspicuous invertebrates in northern wetlands. Although species replacements are often observed along permanence gradients, the underlying causal mechanisms are poorly understood. In this paper, we report on the distributional patterns of caddisflies in permanent and temporary high‐altitude ponds, and how those patterns reflect differences in life history characteristics that affect desiccation tolerance (fundamental niches) versus constraints related to biotic interactions (realised niches). 2. Species (Hesperophylax occidentalis and Agrypnia deflata) that were encountered only in permanent ponds are restricted in distribution by life history (no ovarian diapause, aquatic oviposition, and/or inability to tolerate desiccation). Although the egg masses of H. occidentalis tolerate desiccation, the larvae leave the protective gelatinous matrix of the egg mass because adults oviposit in water. 3. Three species (Asynarchus nigriculus, Limnephilus externus and L. picturatus) have life history characteristics (rapid larval growth, ovarian diapause and terrestrial oviposition of desiccation‐tolerant eggs) that should facilitate the use of both permanent and temporary habitats. However, A. nigriculus is rare or absent in most permanent ponds, and L. externus and L. picturatus are rare or absent in most temporary ponds. Experimental data from a previous study on the combined effects of salamander predation and interspecific interactions among caddisflies (e.g. intraguild predation) suggest that biotic interactions limit each species to a subset of potentially exploitable habitats. 4. Many wetland invertebrates exhibit species replacements along permanence gradients, but few studies have separated the relative importance of the effects of drying per se from the effects of biotic interactions. Our results emphasise the complementary roles of comparative data on life histories and experimental data on competition and predation for understanding invertebrate distributions along permanence gradients.

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 support@jstor.org.. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology. Abstract.Many populations are heterogeneous collections of different sizes or stages of conspecifics. Existing overlap indices do not account for the size-/stage-structured nature of these populations. In this paper I present new overlap indices that use information about the sizes of individuals that co-occur in time and space to predict the potential for interactions in size-structured guilds. An index of the opportunity for competition (JOC) calculates the frequency with which similar size classes of two species encounter each other, whereas an index of the opportunity for intraguild predation (1OP) calculates the frequency of encounters among disparate size classes of the same two species.To illustrate that these indices are more appropriate for size-structured populations than conventional indices, I calculated overlap, 1OP, and IOC for all species pairs in a diverse assemblage of dragonfly larvae. The new indices revealed size-specific patterns of overlap that were not detected by the conventional index, including that (1) some species with high overlap values should interact mainly as competitors, others mainly as intraguild predators, and many as both competitors and predators, (2) subtle differences in phenology and/or size-specific shifts in habitat distribution can lead to the potential for asymmetric interspecific interactions, (3) some species with low pairwise IOP and IOC values are nonetheless vulnerable to the effects of diffuse competition or intraguild predation, (4) seasonal segregation reduces competitive overlap but at the same time increases the opportunity for intraguild predation. The indices are general in form and should be useful for analyzing distributional data for any size-structured assemblage in which the type and intensity of interaction varies asa function of relative size. SV 2 PT 3 4 5 6 7 8 9 HEAD WIDTH (mm) FIG. 1. Interspecific size variation among 12 species of larval dragonflies. Head and gape (labium) widths are for the final instar of each species. Size data for the numerous smaller instars of each species are given in Appendix 1. The abbreviations here are also used in Tables 1-3: PT = Perithemis tenera, LLY = Libellula lydia, LLU = Libellula luctuosa, TL = Tramea lacerate, EC = Epitheca cynosure, ES = Erythemis simplicicollis, SV = 5Sympetrum vicinum, AJ = Anaxjunius, LP = Libellula pulchella, PXL = Pachydiplax longipennis, EP = Epitheca princes, CEL = Celithemis elisa.

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Comparative data from subalpine wetlands in Colorado indicate that larvae of the limnephilid caddisfl.ies, Asynarchus nigriculus and Limnephilus externus, are reciprocally abundant among habitats-Limnephilus larvae dominate in permanent waters, whereas Asynarchus larvae dominate in temporary basins. The purpose of this paper is to report on field and laboratory experiments that link this pattern of abundance to biotic interactions among larvae. In the first field experiment, growth and survival were compared in single and mixed species treatments in littoral enclosures. Larvae, which eat mainly vascular plant detritus, grew at similar rates among treatments in both temporary and permanent habitats suggesting that exploitative competition is not important under natural food levels and caddis fly densities. However, the survival of Limnephilus larvae was reduced in the presence of Asynarchus larvae. Subsequent behavioral studies in laboratory arenas revealed that Asynarchus larvae are extremely aggressive predators on Limnephilus larvae. In a second field experiment we manipulated the relative sizes of larvae and found that Limnephilus larvae were preyed on only when Asynarchus larvae had the same size advantage observed in natural populations. Our data suggest that the dominance of Asynarchus larvae in temporary habitats is due to asymmetric intraguild predation (IGP) facilitated by a phenological head start in development. These data do not explain the dominance of Limnephilus larvae in permanent basins, which we show elsewhere to be an indirect effect of salamander predation.Behavioral observations also revealed that Asynarchus larvae are cannibalistic. In contrast to the IGP on Limnephilus larvae, Asynarchus cannibalism occurs among same-sized larvae and often involves the mobbing of one victim by several conspecifics. In a third field experiment, we found that Asynarchus cannibalism was not density-dependent and occurred even at low larval densities. We hypothesize that Asynarchus IGP and cannibalism provide a dietary supplement to detritus that may be necessary for the timely completion of development in these nutrient-poor, high-elevation wetlands.

We conducted a series of field and laboratory experiments to determine the direct and indirect effects of a top predator, the tiger salamander (Ambystoma tigrinum nebulosum), on larvae of two species of limnephilid caddisflies (Limnephilus externus and Asynarchus nigriculus) in subalpine wetlands in central Colorado. Asynarchus larvae predominate in temporary wetlands and are aggressive intraguild predators on Limnephilus larvae, which only predominate in permanent basins with salamanders. We first conducted a field experiment in mesocosms (cattle tanks) to quantify the predatory effects of different life stages of salamanders on the two caddisfly species. Two life stages of the salamanders (larvae and paedomorphs) preferentially preyed on Asynarchus relative to Limnephilus. Subsequent laboratory experiments revealed that high Asynarchus activity rates and relatively ineffective antipredatory behaviors led to higher salamander detection and attack rates compared to Limnephilus. In a second field experiment (full factorial for presence and absence of each of the three species), we found that salamander predation on Asynarchus had an indirect positive effect on Limnephilus: survival was higher in the presence of salamanders ϩ Asynarchus than with just Asynarchus. In the laboratory we compared the predatory effects of salamanders with and without their mouths sewn shut and found the observed indirect positive effect on Limnephilus survival to be mainly the result of reduced numbers of Asynarchus rather than salamander-induced changes in Asynarchus behavior. We argue that indirect effects of predator-predator interactions on shared prey will be mainly density-mediated and not trait-mediated when one of the predators (in this case, Asynarchus) is under strong selection for rapid growth and therefore does not modify foraging behaviors in response to the other predator. The reciprocal dominance of Limnephilus and Asynarchus in habitats with and without salamanders probably reflects a tradeoff between competitive superiority and vulnerability to predation. The high activity levels and aggressiveness that enable Asynarchus to complete development in temporary habitats result in strong asymmetric competition (via intraguild predation) with Limnephilus. In permanent habitats these same behaviors increase Asynarchus vulnerability to salamander predation, which indirectly benefits Limnephilus. This and previous work implicate salamanders as keystone predators that exert a major influence on the composition of benthic and planktonic assemblages in subalpine wetlands.