Predator size and phenology shape prey survival in temporary ponds

Theoretical efforts suggest that the relative sizes of predators and their prey can shape community dynamics, the structure of food webs, and the evolution of life histories. However, much of this work has assumed static predator and prey body sizes. The timing of recruitment and the growth patterns of both predator and prey have the potential to modify the strength of predator–prey interactions. In this study, I examined how predator size dynamics in 40 temporary ponds over a 3-year period affected the survival of spotted salamander (Ambystoma maculatum) larvae. Across communities, gape-limited predator richness, but not size, was correlated with habitat duration (pond permanence). Within communities, mean gape-limited predator size diminished as the growing season progressed. This size reduction occurred because prey individuals grew into a body size refuge and because the largest of the predators left ponds by mid-season. Elevated gape-limited predation risk across time and space was predicted by the occurrence of two large predatory salamanders: marbled salamander larvae (Ambystoma opacum) and red-spotted newt adults (Notophthalmus viridescens). The presence of the largest gape-limited predator, A. opacum, predicted A. maculatum larval survival in the field. The distribution of large predatory salamanders among ponds and across time is expected to lead to differing community dynamics and to generate divergent natural selection on early growth and body size in A. maculatum. In general, a dynamic perspective on predator size often will be necessary to understand the ecology and evolution of species interactions. This will be especially true in frequently disturbed or seasonal habitats where phenology and ontogeny interact to determine body size asymmetries. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Competition during early ontogeny: Effects of native and invasive planktivores on the growth,survival, and habitat use of bluegill

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The non-consumptive effects of predators and personality on prey growth and mortality

Individual organisms vary in personality, and the ecological consequences of that variation can affect the strength of predator–prey interactions. Prey with bolder tendencies can mitigate the strength of species interactions by altering growth and initiating ontogenetic niche shifts (ONS). While the link between personality and growth has been established, recent research has highlighted the important interplay between ONS and predator cues in community ecology. The objective of this study was to evaluate the effects of prey personality and predator cues on prey growth and ONS. We predicted growth–mortality trade-offs among personalities with higher survival, larger size, and accelerated ONS for bold individuals in comparison with shy individuals. To evaluate this objective, we conducted behavioral assays and a mesocosm experiment to test how southern leopard frog (Rana sphenocephala) tadpole personality and predatory fish (bluegill, Lepomis macrochirus) cues affects tadpole growth and metamorphosis. On average, bold tadpoles had higher mortality across all treatments in comparison with shy tadpoles. The effects of fish cues were dependent on tadpole personality with shy tadpoles metamorphosing significantly later than bold tadpoles. Bold tadpoles were larger than shy tadpoles at metamorphosis; however, that pattern reversed with fish cues as shy individuals metamorphosed larger than bold individuals. Our results suggest personality may be useful for predicting growth and life history for some prey species with predators. Specifically, the threat of predation can interact with personality to incur a benefit (earlier ONS) while also incurring a cost (size at metamorphosis). Hence by incorporating predator cues with personality, ecologists will be able to elucidate growth–mortality trade-offs mediated by personality.     

Dynamics of handling time and functional response by larvae of Chrysomya albiceps (Dipt., Calliphoridae) on different prey species

Abstract:  One way to understand the behavioural patterns exhibited by a predator in response to prey density is to evaluate its functional response. Such evaluation yields information about basic mechanisms of prey–predator dynamics, and is an essential component of prey–predator models. In this paper we analysed experimentally the functional response and the handling time spent by Chrysomya albiceps on different prey species and larval instars of blowflies. The type II functional response was observed when second instar larvae of Chrysomya megacephala and Chrysomya macellaria were consumed. The handling time spent by the predator was significantly different between instars and species. The implications of the functional response and handling time for the interaction dynamics of Brazilian Chrysomyinae species are discussed.     

Effects of food-deprivation on migratory restlessness and diurnal activity in the garden warbler Sylvia borin

Summary The effects of tadpole body size, tadpole sibship, and fish body size on predation of gray treefrog tadpoles, Hyla chrysoscelis, were studied in laboratory and artificial pond experiments. Tadpole body size had a significantly positive effect on the survival of tadpoles in all experiments. The relationship between tadpole biomass eaten and biomass available suggested that fish were not satiated when consuming the largest tadpoles. Large tadpoles were probably better able to evade predators. A difference in survival among full sib families of tadpoles was only present in one family, suggesting that genetic differences in predator avoidance behavior or palatability were probably secondarily important to body size per se. Fish body size had a significantly negative effect on the survival of tadpoles. Larger fish consumed a larger number and proportion of tadpoles as well as greater biomass. These results indicate that environmental factors affecting the growth rate of tadpoles cand dramatically alter their vulnerability to gape-limited predators.     

The foraging ecology of larval and juvenile fishes

Knowledge of the foraging ecology of fishes is fundamental both to understanding the processes that function at the individual, population and community levels, and for the management and conservation of their populations and habitats. Furthermore, the factors that influence the acquisition and assimilation of food can have significant consequences for the condition, growth, survival and recruitment of fishes. The majority of marine and freshwater fish species are planktivorous at the onset of exogenous nutrition and have a limited ability to detect, capture, ingest and digest prey. Improvements in vision, development of fins and associated improvements in swimming performance, increases in gape size and development of the alimentary tract during ontogeny often lead to shifts in diet composition. Prey size, morphology, behaviour and abundance can all influence the prey selection of larval and juvenile fishes. Differences in feeding behaviour between fish species, individuals or during ontogeny can also be important, as can inter- and intraspecific interactions (competition, predation risk). Temporal (diel, seasonal, annual) and spatial (microhabitat, mesohabitat, macrohabitat, regional) variations in prey availability can have important implications for the prey selection, diet composition, growth, survival, condition and, ultimately, recruitment success of fishes. For fish populations to persist, habitat must be available in sufficient quality and quantity for the range of activities undertaken during all periods of development. Habitats that enhance the diversity, size ranges and abundance of zooplankton should ensure that sufficient food resources are available to larval and juvenile fishes.     

Patch size influences perceived patch quality for colonising Culex mosquitoes

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Behavioural and life history effects of predator diet cues during ontogeny in damselfly larvae

A central issue in predator–prey interactions is how predator associated chemical cues affect the behaviour and life history of prey. In this study, we investigated how growth and behaviour during ontogeny of a damselfly larva (Coenagrion hastulatum) in high and low food environments was affected by the diet of a predator (Aeshna juncea). We reared larvae in three different predator treatments; no predator, predator feeding on conspecifics and predator feeding on heterospecifics. We found that, independent of food availability, larvae displayed the strongest anti-predator behaviours where predators consumed prey conspecifics. Interestingly, the effect of predator diet on prey activity was only present early in ontogeny, whereas late in ontogeny no difference in prey activity between treatments could be found. In contrast, the significant effect of predator diet on prey spatial distribution was unaffected by time. Larval size was affected by both food availability and predator diet. Larvae reared in the high food treatment grew larger than larvae in the low food treatment. Mean larval size was smallest in the treatment where predators consumed prey conspecifics, intermediate where predators consumed heterospecifics and largest in the treatment without predators. The difference in mean larval size between treatments is probably an effect of reduced larval feeding, due to behavioural responses to chemical cues associated with predator diet. Our study suggests that anti-predator responses can be specific for certain stages in ontogeny. This finding shows the importance of considering where in its ontogeny a study organism is before results are interpreted and generalisations are made. Furthermore, this finding accentuates the importance of long-term studies and may have implications for how results generated by short-term studies can be used.     

Multi-predator effects across life-history stages: non-additivity of egg- and larval-stage predation in an African treefrog

The effects of multiple predators on their prey are frequently non‐additive because of interactions among predators. When prey shift habitats through ontogeny, many of their predators cannot interact directly. However, predators that occur in different habitats or feed on different prey stages may still interact through indirect effects mediated by prey traits and density. We conducted an experiment to evaluate the combined effects of arboreal egg‐stage and aquatic larval‐stage predators of the African treefrog, Hyperolius spinigularis. Egg and larval predator effects were non‐additive – more Hyperolius survived both predators than predicted from their independent effects. Egg‐stage predator effects on aquatic larval density and size and age at hatching reduced the effectiveness of larval‐stage predators by 70%. Our results indicate that density‐ and trait‐mediated indirect interactions can act across life‐stages and habitats, resulting in non‐additive multi‐predator effects.     

Seasonal Trophic Niche Shift and Cascading Effect of a Generalist Predator Fish

Few studies have examined how foraging niche shift of a predator over time cascade down to local prey communities. Here we examine patterns of temporal foraging niche shifts of a generalist predator (yellow catfish, Pelteobagrus fulvidraco) and the abundance of prey communities in a subtropical lake. We predicted that the nature of these interactions would have implications for patterns in diet shifts and growth of the predator. Our results show significant decreases in planktivory and benthivory from late spring to summer and autumn, whereas piscivory increased significantly from mid-summer until late autumn and also increased steadily with predator body length. The temporal dynamics in predator/prey ratios indicate that the predation pressure on zooplankton and zoobenthos decreased when the predation pressure on the prey fish and shrimps was high. Yellow catfish adjusted their foraging strategies to temporal changes in food availability, which is in agreement with optimal foraging theory. Meanwhile the decrease in planktivory and benthivory of yellow catfish enabled primary consumers, such as zooplankton and benthic invertebrates, to develop under low grazing pressure via trophic cascading effects in the local food web. Thus, yellow catfish shifts its foraging niche to intermediate consumers in the food web to benefit the energetic demand on growth and reproduction during summer, which in turn indirectly facilitate the primary consumers. In complex food webs, trophic interactions are usually expected to reduce the strength and penetrance of trophic cascades. However, our study demonstrates strong associations between foraging niche of piscivorous fish and abundance of prey. This relationship appeared to be an important factor in producing top-down effects on both benthic and planktonic food webs.     

Associations between Non‐Lethal Injury,Body Size,and Foraging Ecology in an Amphibian Intraguild Predator

Theoretical treatments of intraguild predation and its effects on behavioral interactions regard the phenomenon as a size‐structured binary response wherein predation among competitors is completely successful or completely unsuccessful. However, intermediate outcomes occur when individuals escape intraguild (IG) interactions with non‐lethal injuries. While the effects of wounds for prey include compromised mobility and increased predation risk, the consequences of similar injuries among top predators are not well understood, despite the implications for species interactions. Using an amphibian IG predator, Ambystoma opacum (Caudata: Ambystomatidae), we examined associations between non‐lethal injuries and predator body size, foraging strategy, microhabitat selection, and intraspecific agonistic interactions. Wounds were common among IG predators, generally increasing in frequency throughout larval ontogeny. Non‐lethal injuries were associated with differences in predator body size and behavior, with injured predators exhibiting smaller body sizes, increased use of benthic microhabitats, reduced agonistic displays, and increased risk of intraspecific aggression. While such effects were not ultimately associated with reduced foraging success, non‐lethal injury could contribute to niche partitioning between injured and healthy predators via habitat selection, but injured predators likely continue to exert predatory pressure on IG and basal prey populations. Our results indicate that studies of top‐down population regulation should incorporate injury‐related modifications to both prey and predator behavior and size structure.     

Pattern and process in the geographical ranges of freshwater fishes

North American freshwater fishes were studied to determine whether they displayed the same relationships between log (geographical range size) and log (body size) and the same pattern of range shape as found among North American birds and mammals. The forces that produce these patterns were also investigated. The log (geographical range size) : log (body size) relationship was analysed for 121 North American freshwater fish species. Thirty‐two imperilled species were compared with 89 non‐imperilled species to determine if the overall relationship could result from differential extinction. Range geometries were analysed, within and among habitat guilds, to determine if general patterns could be detected. The log (geographical range size) : log (body size) pattern among freshwater fish species was triangular and qualitatively similar to that found for North American birds and mammals. The results suggest that below a minimum geographical range, the likelihood of extinction increases dramatically for freshwater fishes and that this minimum range size increases with body size. The pattern of fish species’ range shapes differs from that found for other North American vertebrate taxa because, on average, fish possess much smaller ranges than terrestrial species and most fish species’ geographical ranges extend further on a north–south axis than on an east–west axis. The log (geographical range size) : log (body size) pattern reveals that fish species’ geographical ranges are more constrained than those of terrestrial species. The triangular relationship may be caused by differential extinction of species with large bodies and small geographical ranges as well as higher speciation rates of small‐bodied fish. The restricted geographical ranges of freshwater fishes gives them much in common with terrestrial species on oceanic islands. Range shape patterns within habitat guilds reflect guild‐specific historical and current ecological forces. The overall pattern of range shapes emerges from the combination of ecologically different subunits.     

Seasonal migration determined by a trade-off between predator avoidance and growth

Migration is a common phenomenon in many organisms, terrestrial as well as aquatic, and considerable effort has been spent to understand the evolution of migratory behaviour and its consequences for population and community dynamics. In aquatic systems, studies on migration have mainly been focused on commercially important fish species, such as salmon and trout. However, seasonal mass-migrations may occur also among other freshwater fish, e.g. in cyprinids that leave lakes and migrate into streams and wetlands in the fall and return back to the lake in spring. In a conceptual model, we hypothesized that this is an adaptive behaviour in response to seasonal changes in predation (P) and growth (G) and that migrating fish change habitat so as to minimise the ratio between predation mortality and growth rate (P/G). Estimates from bioenergetic modelling showed that seasonal changes in the ratio between predator consumption rate and prey growth rate followed the predictions from the conceptual model and also gave more precise predictions for the timing of the habitat change. By quantifying the migration of more than 1800 individually marked fish, we showed that actual migration patterns followed predictions with a remarkable accuracy, suggesting that migration patterns have evolved in response to seasonally fluctuating trade-offs between predator avoidance and foraging gains. Thus, the conceptual model provides a mechanistic understanding to mass-migration in prey fish. Further, we also show that the dominant prey fish is actually absent from the lake during a major part of the year, which should have strong implications for the dynamics of the lake ecosystem through direct and indirect food-web interactions.     

Effects of avian predation threat, water flow and cover on growth and habitat use by chub, Leuciscus cephalus, in an experimental stream

Trophic interactions, including “top‐down” predator‐prey interactions, are particularly important in influencing the structure of fish communities. While the varied impacts of piscivorous fish have been well investigated, the effects of fish‐eating birds on riverine fish behaviour and population dynamics still remain controversial, mainly because they are undervalued. Summer experiments were conducted in an experimental outdoor stream to evaluate the effects of avian predation threat, stream flow, and overhead cover on growth and behavioural tactics of wild juvenile chub (Leuciscus cephalus). Groups of fifteen chub maintained in riffle‐pool sequences were submitted to combinations of different conditions, namely absence or presence of a simulated fish‐eating bird, low or high flow, and absence or presence of medium or high cover. In the absence of predation threat, chub foraged in the riffles and maximized feeding opportunities. Under predation threat, they sheltered, foraged less and grew slowly and as expected, they increased their use of the riffles at high flow as water turbulence is an efficient shelter from birds but only in the absence of cover. In the presence of cover, fish sheltered exclusively under pool covers and were more prone to take risks at low flow because of higher costs in terms of lost feeding opportunities associated with these conditions. This result indicates that flow velocity altered cover use tactics through its impact on food supply, suggesting that it may affect the outcome of predator‐prey relationships. So, chub use cover in a flexible manner according to both the benefits in terms of predator avoidance and the costs in terms of lost feeding opportunities. A striking finding of the experiments is the drastic reduction in the range of growth variances amongst fish when they are maintained under predation threat, suggesting a homogenization of fitness between individuals. From all our results, we argue that in lowland streams, under summer field situations, fish‐eating birds may affect local prey population dynamics more through sub‐lethal effects on growth rates than directly through death rates.     

And the Last Shall Be First: Heterochrony and Compensatory Marine Growth in Sea Trout (Salmo trutta)

Early juvenile growth is a good indicator of growth later in life in many species because larger than average juveniles tend to have a competitive advantage. However, for migratory species the relationship between juvenile and adult growth remains obscure. We used scale analysis to reconstruct growth trajectories of migratory sea trout (Salmo trutta) from six neighbouring populations, and compared the size individuals attained in freshwater (before migration) with their subsequent growth at sea (after migration). We also calculated the coefficient of variation (CV) to examine how much body size varied across populations and life stages. Specifically, we tested the hypothesis that the CV on body size would differ between freshwater and marine environment, perhaps reflecting different trade-offs during ontogeny. Neighbouring sea trout populations differed significantly in time spent at sea and in age-adjusted size of returning adults, but not on size of seaward migration, which was surprisingly uniform and may be indicative of strong selection pressures. The CV on body size decreased significantly over time and was highest during the first 8 months of life (when juvenile mortality is highest) and lowest during the marine phase. Size attained in freshwater was negatively related to growth during the first marine growing season, suggesting the existence of compensatory growth, whereby individuals that grow poorly in freshwater are able to catch up later at sea. Analysis of 61 datasets indicates that negative or no associations between pre- and post-migratory growth are common amongst migratory salmonids. We suggest that despite a widespread selective advantage of large body size in freshwater, freshwater growth is a poor predictor of final body size amongst migratory fish because selection may favour growth heterochrony during transitions to a novel environment, and marine compensatory growth may negate any initial size advantage acquired in freshwater.     

Feedback between size balance and consumption strongly affects the consequences of hatching phenology in size‐dependent predator–prey interactions

In many size‐dependent predator–prey systems, hatching phenology strongly affects predator–prey interaction outcomes. Early‐hatched predators can easily consume prey when they first interact because they encounter smaller prey. However, this process by itself may be insufficient to explain all predator–prey interaction outcomes over the whole interaction period because the predator–prey size balance changes dynamically throughout their ontogeny. We hypothesized that hatching phenology influences predator–prey interactions via a feedback mechanism between the predator–prey size balance and prey consumption by predators. We experimentally tested this hypothesis in an amphibian predator–prey model system. Frog tadpoles Rana pirica were exposed to a predatory salamander larva Hynobius retardatus that had hatched 5, 12, 19 or 26 days after the frog tadpoles hatched. We investigated how the salamander hatch timing affected the dynamics of prey mortality, size changes of both predator and prey, and their subsequent life history (larval period and size at metamorphosis). The predator–prey size balance favoured earlier hatched salamanders, which just after hatching could successfully consume more frog tadpoles than later hatched salamanders. The early‐hatched salamanders grew rapidly and their accelerated growth enabled them to maintain the predator‐superior size balance; thus, they continued to exert strong predation pressure on the frog tadpoles in the subsequent period. Furthermore, frog tadpoles exposed to the early‐hatched salamanders were larger at metamorphosis and had a longer larval period than other frog tadpoles. These results suggest that feedback between the predator‐superior size balance and prey consumption is a critical mechanism that strongly affects the impacts of early hatching of predators in the short‐term population dynamics and life history of the prey. Because consumption of large nutrient‐rich prey items supports the growth of predators, a similar feedback mechanism may be common and have strong impacts on phenological shifts in size‐dependent trophic relationships.     

The non-lethal effects of predators and the influence of food availability on life history of adult Chironomus tentans (Diptera: Chironomidae)

1. We used a laboratory experiment to determine effects of a predator (other than straightforward prey consumption) and food availability, during the larval stage, on adult size, age at emergence and fecundity of Chironomus tentans. 2. Predator presence and decreased food availability resulted in reduced adult emergence. 3. Predator presence and food availability resulted in smaller size and greater age at emergence of male and female chironomids. There were no significant interactions between effects of predator presence and food availability. 4. Predator presence had no significant effect on fecundity. 5. Smaller size and greater age at emergence can have important implications for adult survival, size and age at first reproduction and, thus, intrinsic rate of population growth.     

Interactive effects of habitat selection, food supply and predation on recruitment of an estuarine fish

 Seagrass meadows are often important habitats for newly recruited juvenile fishes. Although substantial effort has gone into documenting patterns of association of fishes with attributes of seagrass beds, experimental investigations of why fish use seagrass habitats are rare. We performed two short-term manipulative field experiments to test (1) the effects of food supply on growth and densities of fish, and (2) effects of predation on the density and size distribution of fish recruits, and how this varies among habitat types. Experiments were conducted in Galveston Bay, Texas, and we focused on the common estuarine fish, pinfish Lagodon rhomboides. In the first experiment, replicate artifical seagrass and sand plots were either supplemented with food or left as controls. Recruitment of pinfish was significantly greater to seagrass than sand habitats; however, we detected no effect of food supplementation on the abundance of recruits in either habitat. Pinfish recruits in artifical seagrass grew at a significantly faster rate than those in sand habitats, and fish supplemented with food exhibited a greater growth rate than controls in both sand and artifical grass habitats. In our second experiment, we provided artificial seagrass and sand habitats with and without predator access. Predator access was manipulated with cages, and two-sided cages served as controls. Recruitment was significantly greater to the cage versus cage-control treatment, and this effect did not vary between habitats. In addition, the standard length of pinfish recruits was significantly larger in the predator access than in the predator exclusion treatment, suggesting size-selective predation on smaller settlers or density-dependent growth. Our results indicate that the impact of predation on pinfish recruits is equivalent in both sand and vegetated habitats, and thus differential predation does not explain the higher recruitment of pinfish to vegetated than to nonvegetated habitats. Since predators may disproportionately affect smaller fish, and a limited food resource appears to be more effectively utilized by fish in vegetated than in unvegetated habitats, we hypothesize that pinfish recruits may select vegetated habitats because high growth rates allow them to achieve a size that is relatively safe from predation more quickly. Received: 10 October 1996 / Accepted: 5 April 1997     

Mechanisms influencing competition between hatchery and wild juvenile anadromous Pacific salmonids in fresh water and their relative competitive abilities

Avoiding negative effects of competition from released hatchery salmonids on wild fish is a primary concern for recovery efforts and fisheries management. Several factors affect competition among juvenile salmonids including: (1) whether competition is intra- or interspecific, (2) duration of freshwater cohabitation of hatchery and wild fish, (3) relative body size, (4) prior residence, (5) environmentally induced developmental differences, and (6) fish density. Intraspecific competition is expected to be greater than interspecific because of greater niche overlap between conspecific hatchery and wild fish. Competition is expected to increase with prolonged freshwater cohabitation. Hatchery smolts are often larger than wild, and larger fish are usually superior competitors. However, wild fish have the advantage of prior residence when defending territories and resources in natural streams. Hatchery-induced developmental differences are variable and can favor both hatchery and wild fish. Although all these factors influence competitive interactions, fish density of the composite population (wild + hatchery fish) in relation to habitat carrying capacity likely exerts the greatest influence. The extent of competition and relative competitive ability of wild and hatchery fish can be determined by additive and substitutive experimental designs, respectively, and the limited body of substitutive experiments suggests that the relative competitive ability of hatchery and wild fish is approximately equal when measured as growth. Conducting substitutive experiments becomes difficult as the spatial and temporal scales increase. Large-scale experiments comparing supplemented and control reaches or streams hold some promise for quantifying the effects of released hatchery fish on wild fish behavior, growth and survival.     

Well-informed foraging: damage-released chemical cues of injured prey signal quality and size to predators

Predators use a variety of information sources to locate potential prey, and likewise prey animals use numerous sources of information to detect and avoid becoming the meal of a potential predator. In freshwater environments, chemosensory cues often play a crucial role in such predator/prey interactions. The importance of chemosensory information to teleost fish in marine environments is not well understood. Here, we tested whether coral reef fish predators are attracted to damage-released chemical cues from already wounded prey in order to find patches of prey and minimize their own costs of obtaining food. Furthermore, we tested if these chemical cues would convey information about status of the prey. Using y-maze experiments, we found that predatory dottybacks, Pseudochromis fuscus, were more attracted to skin extracts of damselfish, Pomacentrus amboinensis, prey that were in good condition compared to prey in poor body condition. Moreover, in both the laboratory and field, we found that predators could differentiate between skin extracts from prey based on prey size, showing a greater attraction to extracts made from prey that were the appropriate size to consume. This suggests that predators are not attracted to any general substance released from an injured prey fish instead being capable of detecting and distinguishing relatively small differences in the chemical composition of the skin of their prey. These results have implications for understanding predator foraging strategies and highlights that chemical cues play a complex role in predator–prey interactions in marine fish.