Scott A. Wissinger

Niche Overlap and the Potential for Competition and Intraguild Predation Between Size‐Structured Populations

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 (IOC) calculates the frequency with which similar size classes of two species encounter each other, whereas an index of the opportunity for intraguild predation (IOP) 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, IOP, 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 as a function of relative size.

Foraging trade-offs along a predator-permanence gradient in subalpine 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 pre- dominate 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 rela- tively 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 + 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 Lim- nephilus and Asynarchus in habitats with and without salamanders probably reflects a trade- off 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 sala- manders as keystone predators that exert a major influence on the composition of benthic and planktonic assemblages in subalpine wetlands.

Intraguild Predation and Cannibalism among Larvae of Detritivorous Caddisflies in Subalpine Wetlands

Comparative data from subalpine wetlands in Colorado indicate that larvae of the limnephilid caddisflies, Asynarchus nigriculus and Limnephilus externus, are recip- rocally 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 caddisfly 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 ad- vantage 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 con- trast 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.

Growth and foraging consequences of facultative paedomorphosis in the tiger salamander,Ambystoma tigrinum nebulosum

SummaryFacultative paedomorphosis in salamanders occurs when larvae respond to varying environmental conditions by either metamorphosing into terrestrial metamorphic adults or retaining their larval morphology to become sexually mature paedomorphic adults. Several hypotheses have been proposed for the evolutionary maintenance of this environmentally induced dimorphism, but few data are available to assess them adequately. We studied a montane population of the tiger salamander,Ambystoma tigrinum nebulosum, and measured the adult growth rate and body condition across three growing seasons to assess the relative costs and benefits of each morph. Metamorphic adults grew more than paedomorphic adults in terms of snout—vent length across years and in weight within years. Dietary analyses and foraging experiments revealed some of the proximate factors that may underlie these differential growth patterns. Across all prey, metamorphs had significantly higher biomass and calories per stomach sample than paedomorphs. Metamorphic diets primarily consisted of the fairy shrimpBranchinecta coloradensis, whereas paedomorphic diets contained a variety of benthic and terrestrial invertebrates. Foraging experiments revealed that both morphs are more successful at capturing fairy shrimp relative to other prey types and both show high electivity toward this prey. However, fairy shrimp occurred only in non-permanent ponds and thus are inaccessible to paedomorphs, which can survive only in permanent ponds. Paedomorphs also experience higher levels of intraspecific competition with large larvae in permanent ponds than metamorphs do in non-permanent ponds. Thus, metamorphs obtain a growth advantage over paedomorphs by foraging in non-permanent ponds that contain fairy shrimp and have reduced intraspecific competition. These results suggest that paedomorphs should have decreased fitness relative to metamorphs, primarily because metamorphs can move into the best habitats for growth. The net fitness effect of morph-specific differences in dispersal depend on whether there are trade-offs with other life history traits. Nonetheless, because the relative benefit of metamorph dispersal will change with environmental conditions in permanent ponds and the surrounding habitat, the relative fitness payoff to each morph should track changes in the environment. Thus, facultative paedomorphosis may be maintained in part by variable, environmentally-specific fitness payoffs to each morph.

Life history and size structure of larval dragonfly populations

Comparative data on emergence, adult activity, and larval development were used to interpret patterns of phenology, voltinism, and developmental synchrony in a 14-species assemblage of dragonflies (Odonata:Anisoptera). Interspecific differences in these characteristics provided an opportunity to interrelate life history, population size structure, and the potential for intra- and interspecific interactions. Larval populations of most species comprised an extraordinary size range of individuals at any one time. In many species, this intraspecific size variability was primarily due to developmental asynchrony within generations (e.g., Libellula lydia, Celithemis elisa, Perithemis tenera). Oviposition in these species occurred throughout most of the summer so that the timing of instar development varied considerably among broods. By fall, the largest larvae in these populations were 5 to 10 times the size of smaller conspecifics. In other asynchronous breeders, an even wider size range of co-occurring conspecifics resulted from overlap between generations. In Erythemis simplicicollis and Pachydiplax longipennis this generation overlap was due to the presence of mixed uni- and bivoltine cohorts, whereas in Tramea lacerata and Anax junius, it was due to phenologically separate but overlapping migratory and resident cohorts. In the semivoltine Epitheca princeps, size variability among conspecifics was due mainly to overlap between year classes. Most larvae were similar in size at the same point in time in those species that bred synchronously and were univoltine (Epitheca cynosura, Leucorrhinia intacta). Size ratios of the smallest to largest co-occurring conspecifics in these populations were frequently larger than those commonly thought to facilitate interspecific coexistence. Thus, intraspecific resource partitioning should play an important role in population regulation, as should cannibalism. This intraspecific size variability should also create the potential for mixed competition--predation interactions between the same species. A phenological tradeoff between competition and predation should minimize the efficacy of seaonal segregation as a mechanism for minimizing the intensity of interspecific interaction in this assemblage.

Absence of species replacements between permanent and temporary lentic communities in New Zealand

Abstract The species composition of lentic communities often shifts along hydroperiod gradients, in part because temporary-habitat specialists replace closely related permanent-habitat specialists. These replacements reflect tradeoffs between traits that facilitate coexistence with permanent-habitat predators and those that prevent desiccation. The evidence for species replacements and the underlying tradeoffs is considerable in North America, but few studies have explored this pattern in other regions. We compared benthic communities in permanent and temporary habitats on the South Island of New Zealand. Ordination across 58 sites showed that community composition was distinctly different between the 2 types of habitats. Assemblages in permanent habitats had >2× the number of species as those in temporary habitats. We found little evidence for temporary-habitat specialists; i.e., species in temporary communities were a nested subset of those in permanent communities. Quantitative sampling at 12 intensively studied sites revealed that chironomids, water bugs, beetles, and crustaceans accounted for 90% of the biomass in temporary, but only 14% of the biomass in permanent habitats, which were dominated by mollusks, annelids, caddisflies, and odonates. Damselflies, dragonflies, caddisflies, and several other large-bodied taxa common in permanent habitats were absent from most temporary habitats. We propose 2 explanations for the absence of species replacements in these groups in the New Zealand habitats that we studied. First, drying is unpredictable within and between years, perhaps precluding the evolution of temporary-habitat specialization. Second, fish predation on benthic invertebrates, a driver for phylogenetic diversification in North America, appears to be comparatively weak in New Zealand. Comparative studies across a range of climates and faunas will be needed to identify the ecological and phylogenetic contexts that favor evolution of generalists vs specialists along permanence gradients.

Consumptive and nonconsumptive effects of cannibalism in fluctuating age‐structured populations

Theory and empirical studies suggest that cannibalism in age-structured populations can regulate recruitment depending on the intensity of intraspecific competition between cannibals and victims and the nature of the cannibalism window, i.e., which size classes interact as cannibals and victims. Here we report on a series of experiments that quantify that window for age-structured populations of salamander larvae and paedomorphic adults. We determined body size limits on cannibalism in microcosms and then the consumptive and nonconsumptive (injuries, foraging and activity, diet, growth) effects on victims in mesocosms with seminatural levels of habitat complexity and alternative prey. We found that cannibalism by the largest size classes (paedomorphs and > or = age 3+ yr larvae) occurs mainly on young-of-the-year (YOY) victims. Surviving YOY and other small larvae had increased injuries, reduced activity levels, and reduced growth rates in the presence of cannibals. Data on YOY survival in an experiment in which we manipulated the density of paedomorphs combined with historical data on the number of cannibals in natural populations indicate that dominant cohorts of paedomorphs can cause observed recruitment failures. Dietary data indicate that ontogenetic shifts in diet should preclude strong intraspecific competition between YOY and cannibals in this species. Thus our results are consistent with previous empirical and theoretical work that suggests that recruitment regulation by cannibalism is most likely when YOY are vulnerable to cannibalism but have low dietary overlap with cannibals. Understanding the role of cannibalism in regulating recruitment in salamander populations is timely, given the widespread occurrences of amphibian decline. Previous studies have focused on extrinsic (including anthropogenic) factors that affect amphibian population dynamics, whereas the data presented here combined with long-term field observations suggest the potential for intrinsically driven population cycles.

Fluctuation in a rocky mountain population of salamanders : anthropogenic acidification or natural variation?

We monitored the demographics of the salamander Ambystoma tigrinum nebulosum as part of a community-wide study on the effects of acidification in sub-alpine (elevation 3600 m) ponds in central Colorado. A decline in A. t. nebulosum at this site from 1982 to 1988 has been hypothesized to result from embryonic mortality during a pulse of acidity that accompanies snowmelt in spring. Since 1988 we have monitored salamander population size, reproduction, and recruitment, and compared survival and individual growth rates among ponds which differ five-fold in acid-neutralizing capacity (ANC)

Larval growth in polyphenic salamanders: making the best of a bad lot

Polyphenisms are excellent models for studying phenotypic variation, yet few studies have focused on natural populations. Facultative paedomorphosis is a polyphenism in which salamanders either metamorphose or retain their larval morphology and eventually become paedomorphic. Paedomorphosis can result from selection for capitalizing on favorable aquatic habitats (paedomorph advantage), but could also be a default strategy under poor aquatic conditions (best of a bad lot). We tested these alternatives by quantifying how the developmental environment influences the ontogeny of wild Arizona tiger salamanders (Ambystoma tigrinum nebulosum). Most paedomorphs in our study population arose from slow-growing larvae that developed under high density and size-structured conditions (best of a bad lot), although a few faster-growing larvae also became paedomorphic (paedomorph advantage). Males were more likely to become paedomorphs than females and did so under a greater range of body sizes than females, signifying a critical role for gender in this polyphenism. Our results emphasize that the same phenotype can be adaptive under different environmental and genetic contexts and that studies of phenotypic variation should consider multiple mechanisms of morph production.

Accuracy Assessment of Skeletochronology in the Arizona Tiger Salamander (Ambystoma Tigrinum Nebulosum)

Abstract Skeletochronology is the most commonly used tool for evaluating the age of amphibians and reptiles. However, the accuracy of skeletochronology is questionable because few studies have utilized individuals of known age to calibrate their results. In this study, the accuracy of skeletochronology was assessed using individual Arizona Tiger Salamanders (Ambystoma tigrinum nebulosum) that were of known ages that varied from 2–15 years. Cross-sections of salamander toes were analyzed to compare the number of LAG (lines of arrested growth) to the actual age of known cohorts. Our results suggest that skeletochronology was an inaccurate tool for estimating age in this population of A. tigrinum nebulosum because it grossly underestimated age. The high intensity of endosteal resorption coupled with rapprochement of the most peripheral LAG may have produced these results. Our research indicates that skeletochronological studies that do not use individuals of known age for calibration may underestimate age.