The impact of larval predators and competitors on the morphology and fitness of juvenile treefrogs

Studies of phenotypic plasticity typically focus on traits in single ontogenetic stages. However, plastic responses can be induced in multiple ontogenetic stages and traits induced early in ontogeny may have lasting effects. We examined how gray treefrog larvae altered their morphology in four different larval environments and whether different larval environments affected the survival, growth, development, and morphology of juvenile frogs at metamorphosis. We then reared these juveniles in terrestrial environments under high and low intraspecific competition to determine whether the initial differences in traits at metamorphosis affected subsequent survival and growth, whether the initial phenotypic differences converged over time, and whether competition in the terrestrial environment induced further phenotypic changes. Larval and juvenile environments both affected treefrog traits. Larval predators induced relatively deep tail fins and short bodies, but there was no impact on larval development. In contrast, larval competitors induced relatively short tails and long bodies, reduced larval growth, and slowed larval development. At metamorphosis, larval predators had no effect on juvenile growth or relative morphology while larval competitors produced juveniles that were smaller and possessed relatively shorter limbs and shorter bodies. After 1 month of terrestrial competition among the juvenile frogs, the initial differences in juvenile morphology did not converge. There were no differences in growth due to larval treatment but there were differences in survival. Individuals that experienced low competition as tadpoles experienced near perfect survival as juvenile frogs but individuals that experienced high competition as tadpoles suffered an 18% decrease in survival as juvenile frogs. There were also morphological responses to juvenile competition, but these changes appear to be due, at least in part, to allometric effects. Collectively, these results demonstrate that larval environments can have profound impacts on the traits and fitness of organisms later in ontogeny.     

Reproductive strategy and carry-over effects for species with complex life histories

We investigated the influences of natal-pool and metamorph characteristics on juvenile survival, age-specific breeding probabilities, and dispersal of wood frogs (Lithobates sylvatica) and used this information to infer how life history strategies of short-lived species may offset risks associated with breeding in highly variable habitats. We used multistate mark-recapture data from eight wood frog populations in Maryland, USA, to investigate the influences of natal-pond and metamorph characteristics on post-metamorphic demographics. We found that post-metamorphic juvenile survival was highly variable and negatively influenced by abiotic conditions experienced during development but showed little relationship to larval density or size at metamorphosis. Estimates of recruitment and dispersal probabilities indicated that males mature earlier than females, and a small percentage of each sex disperse to non-natal pools. Survival probabilities for adults during the non-breeding season were less variable than juvenile rates, lower for females, and negatively related to mean monthly precipitation. Survival of adults during the breeding season was generally very high. We provide the first robust estimates of post-metamorphic vital rates of wood frogs that allow for variation in capture probabilities. We found little evidence for an effect of metamorph traits on juvenile survival, suggesting that wood frogs may be able to overcome initial disadvantages to have similar post-metamorphic performance. Our study suggests that variation in the age of maturity for wood frogs may mitigate risks associated with breeding in a highly variable habitat to maximize lifetime fitness without increasing lifespan, and this strategy is minimally affected by carry-over effects from the larval stage.     

Past and Present Risk: Exposure to Predator Chemical Cues Before and after Metamorphosis Influences Juvenile Wood Frog Behavior

Animals are exposed to different predators over their lifespan. This raises the question of whether exposure to predation risk in an early life stage affects the response to predators in subsequent life stages. In this study, we used wood frogs (Rana sylvatica) to test whether exposure to cues indicating predation risk from dragonfly larvae during the wood frog larval stage affected post‐metamorphic activity level and avoidance of garter snake chemical cues. Dragonfly larvae prey upon wood frogs only during the larval stage, whereas garter snakes prey upon wood frogs during both the larval stage and the post‐metamorphic stage. Exposure to predation risk from dragonflies during the larval stage caused post‐metamorphic wood frog juveniles to have greater terrestrial activity than juvenile wood frogs that were not exposed to larval‐stage predation risk from dragonflies. However, exposure to predation risk as larvae did not affect juvenile wood frog responses to chemical cues from garter snakes. Wood frogs exposed as larvae to predation risk from dragonfly larvae avoided garter snake chemical cues to the same extent as wood frog larvae not exposed to predation risk from dragonfly larvae. Our results demonstrate that while some general behaviors exhibit carry‐over effects from earlier life stages, behavioral responses to predators may remain independent of conditions experienced in earlier life stages.     

Post-metamorphic change in activity metabolism of anurans in relation to life history

Summary Newly-metamorphosed individuals of some species of frogs and toads differ from adults in behavior, ecology, and physiology. These differences may be related to broader patterns of the life histories of different species of frogs. In particular, the length of larval life and the size of a frog at metamorphosis appear to be significant factors in post-metamorphic ontogenetic change. These changes in performance are associated with rapid post-metamorphic increases in oxygen transport capacity. Bufo americanus (American toads) and Rana sylvatica (wood frogs) spend only 2–3 months as tadpoles and metamorphose at body masses of 0.25 g or less. Individuals of these species improve endurance and aerobic capacity rapidly during the predispersal period immediately following metamorphosis. Increases in hematocrit, hemoglobin concentration, and heart mass relative to body mass are associated with this improvement in organismal performance. Rana clamitans (green frogs) spend from 3 to 10 months as larvae and weigh 3 g at metamorphosis. Green frogs did not show immediate post-metamorphic increases in performance. Rana palustris (pickerel frogs) are intermediate to wood frogs and green frogs in length of larval life and in size at metamorphosis, and they are intermediate also in their post-metamorphic physiological changes.American toads and wood frogs appear to delay dispersal from their natal ponds while they undergo rapid post-metamorphic growth and development, whereas green frogs disperse as soon as they leave the water, even before they have fully absorbed their tails. The very small body sizes of newly metamorphosed toads and wood frogs appear to limit the scope of their behaviors. The brief larval periods of these species permit them to exploit transient aquatic habitats, but impose costs in the form of a period of post-metamorphic life in which their activities are restricted in time and space compared to those of adults.     

Interacting effects of predation risk and food availability on larval anuran behaviour and development

Age and size at metamorphosis are two important fitness components in species with complex life cycles. In anurans, metamorphic traits show remarkable phenotypic plasticity, especially in response to changes in growth conditions. It is also possible that the perception of risk directly determines changes in larval period and the size of metamorphs. This study examines how the perception of predation risk affects the timing of and size at metamorphosis in common frogs (Rana temporaria). I raised tadpoles at two risk levels (fish-conditioned water or unconditioned water) crossed with the availability or lack of food at night (all tadpoles had food available in the day). Tadpoles reacted to chemical cues from predatory fish by decreasing activity. A novel behavioural result was a predation×food interaction effect on refuge use, which also accounted for most of the predator main effect: predation risk only caused increased refuge use in the night-starved treatment. Despite these behavioural modifications, the perception of predation risk did not affect growth rate and mass at metamorphosis in a simple way: the effects of food regime on growth and size at metamorphosis were dependent on the level of predation risk as revealed by significant predation×food interaction effects. Tadpoles who had food withheld at night metamorphosed at the smallest size, suggesting a negative relationship between size at metamorphosis and refuge use. Tadpoles raised in fish-conditioned water had longer larval periods than those in unconditioned water, but these differences were significant only if food was available at night. These results conflict with the hypotheses that tadpoles should reduce their larval period or growth rates (and hence metamorphose at a smaller size) as the risk of predation increases. In contrast to predation risk, food availability strongly affected the length of the larval period: night-starved tadpoles metamorphosed relatively early with or without fish stimulus. Thus, early metamorphosis resulted from periods of low food availability, but not from a heightened ”perceived risk” of predation. This example counters the hypothesis of acceleration of the developmental rate (which shortens the time to metamorphosis) as a mechanism to escape a risky environment. Received: 18 August 1999 / Accepted: 10 January 2000     

Carry-over effects of the larval environment on post-metamorphic performance in two hylid frogs

Life history theory and empirical studies suggest that large size or earlier metamorphosis are suitable proxies for increased lifetime fitness. Thus, across a gradient of larval habitat quality, individuals with similar phenotypes for these traits should exhibit similar post-metamorphic performance. Here we examine this paradigm by testing for differences in post-metamorphic growth and survival independent of metamorphic size in a temperate (spring peeper, Pseudacris crucifer) and tropical (red-eyed treefrog, Agalychnis callidryas) anuran reared under differing larval conditions. For spring peepers, increased food in the larval environment increased post-metamorphic growth efficiency more than predicted by metamorphic phenotype and led to increased mass. Similarly, red-eyed treefrogs reared at low larval density ended the experiment at a higher mass than predicted by metamorphic phenotype. These results show that larval environments can have delayed effects not captured by examining only metamorphic phenotype. These delayed effects for the larval environment link larval and juvenile life history stages and could be important in the population dynamics of organisms with complex life cycles.     

Plasticity in newt metamorphosis: the effect of predation at embryonic and larval stages

1. Some organisms under variable predator pressure show induced antipredator defences, whose development incurs costs and may be associated with changes to later performance. This may be of especial relevance to animals with complex life histories involving metamorphosis. 2. This study examines the effect of predation environment, experienced both during embryonic and larval stages, on palmate newt (Triturus helveticus) metamorphosis. Newt eggs were raised until hatching with or without exposure to chemical cues from brown trout (Salmo trutta), and larval development was monitored in the presence or absence of the cues. 3. Exposure to predator cues during the embryonic stage resulted in higher growth rates at the larval stage, reduced time to metamorphosis and size at metamorphosis. Metamorphs also had narrower heads and shorter forelimbs than those from predator‐free treatments. In contrast, exposure to predator cues during the larval stage did not affect metamorph characteristics. 4. These results indicate that developing embryos are sensitive to predator chemical cues and that the responses can extend to later stages. Reversion of induced defences when predation risk ceased was not detected. We discuss the possible adaptive significance of these responses.     

What keeps insects small?—Size dependent predation on two species of butterfly larvae

Insect size usually increases greatly in the latter stages of development, while reproductive value increases strongly with adult size. Mechanisms that can balance the benefits associated with increased growth are poorly understood, raising the question: what keeps insects from becoming larger? If predation risk was to increase with juvenile size, it would make an extension of development very risky, favouring smaller final sizes. But field measures of juvenile mortality seldom show any general patterns of size dependence. We here therefore try to estimate a mechanistic relationship between juvenile size and predation risk by exposing the larvae of two closely related butterflies to a generalist invertebrate predator in a laboratory experiment. Predation risk increased with larval size but was not affected by the species-specific growth rate differences. These results indicate that predation risk may increase with the size of the juvenile even when predators are relatively small. By basing a model simulation on our data we also show that size dependent predation of the kind found in this study has potential to stabilise selection on body size in these species. Thus, these findings suggest that more detailed studies of the size dependence of predation risk on juvenile instars will increase the understanding of what it is that keeps insects small.     

Non-equivalence of growth arrest induced by predation risk or food limitation: context-dependent compensatory growth in anuran tadpoles

1. To gain insight into the evolution of compensatory growth, we studied the growth patterns of anuran (Rana temporaria) larvae following either a period of exogenous growth depression (food restriction) or a period of endogenous depression (exposure to predators). We also investigated the potential deferred costs that larval compensatory growth could impose on post-metamorphic individuals. 2. Food-deprived larvae exhibited full compensatory growth in response to reduced growth rates caused by food limitation, and the growth trajectories of low- and high-rations tadpoles converged before the onset of metamorphosis. 3. According to our predictions, individuals exposed to larval predators did not show growth compensation following predator removal despite undergoing a significant reduction in growth rate associated with low activity levels. 4. Jumping ability of individuals exposed to predators during only 20 days from the commencement of the larval phase was equivalent to that of non-exposed animals, and greater than the jumping capacity of those maintained with predators until the time of metamorphosis. This pattern was consistent with the pattern observed for variation in relative leg length. 5. These results support the suggestion that submaximum and compensatory growth could have evolved to minimize the overall growth/mortality costs in environments with high spatiotemporal variation in predation intensity.     

Behavioral plasticity mitigates risk across environments and predators during anuran metamorphosis

Most animals metamorphose, changing morphology, physiology, behavior and ecological interactions. Size- and habitat-dependent mortality risk is thought to affect the evolution and plastic expression of metamorphic timing, and high predation during the morphological transition is posited as a critical selective force shaping complex life cycles. Nonetheless, empirical data on how risk changes across metamorphosis and stage-specific habitats, or how that varies with size, are rare. We examined predator–prey interactions of red-eyed treefrogs, Agalychnis callidryas, with an aquatic predator (giant water bug, Belostoma) and a semi-terrestrial predator (fishing spider, Thaumasia) across metamorphosis. We manipulated tadpole density to generate variation in metamorph size and conducted predation trials at multiple developmental stages. We quantified how frog behavior (activity) changes across metamorphic development, habitats, and predator presence or absence. In aquatic trials with water bugs, frog mortality increased with forelimb emergence, as hypothesized. In semi-terrestrial trials, contrary to predictions, predation by spiders increased, not decreased, with tail resorption. In neither case did frog size affect mortality. Frogs reduced activity upon forelimb emergence in the water, and further with emergence into air, then increased activity with tail resorption. Longer-tailed metamorphs were captured more often in spider attacks, but attacked less, as most attacks followed prey movements. Metamorphs behaviorally compensated for poor escape performance more effectively on land than in water, thus emergence timing may critically affect mortality. The developmental timing of the ecological transition between environments that select for different larval and juvenile phenotypes is an important, neglected variable in studies of complex life cycles.     

Effects of predation environment and food availability on somatic growth in the Livebearing Fish Brachyrhaphis rhabdophora (Pisces: Poeciliidae)

Variation in somatic growth rates is of great interest to biologists because of the relationship between growth and other fitness‐determining traits, and it results from both genetic and environmentally induced variation (i.e. plasticity). Theoretical predictions suggest that mean somatic growth rates and the shape of the reaction norm for growth can be influenced by variation in predator‐induced mortality rates. Few studies have focused on variation in reaction norms for growth in response to resource availability between high‐predation and low‐predation environments. We used juvenile Brachyrhaphis rhabdophora from high‐predation and low‐predation environments to test for variation in mean growth rates and for variation in reaction norms for growth at two levels of food availability in a common‐environment experiment. To test for variation in growth rates in the field, we compared somatic growth rates in juveniles in high‐predation and low‐predation environments. In the common‐environment experiment, mean growth rates did not differ between fish from differing predation environments, but the interaction between predation environment and food level took the form of a crossing reaction norm for both growth in length and mass. Fish from low‐predation environments exhibited no significant difference in growth rate between high and low food treatments. In contrast, fish from high‐predation environments exhibited variation in growth rates between high and low food treatments, with higher food availability resulting in higher growth rates. In the field, individuals in the high‐predation environment grow at a faster rate than those in low‐predation environments at the smallest sizes (comparable to sizes in the common‐environment experiment). These data provide no evidence for evolved differences in mean growth rates between predation environments. However, fish from high‐predation environments exhibited greater plasticity in growth rates in response to resource availability suggesting that predation environments may exhibit increased variation in food availability for prey fish and consequent selection for plasticity.     

Sequential predator effects across three life stages of the African tree frog, <Emphasis Type="Italic">Hyperolius spinigularis</Emphasis>

While theoretical studies of the timing of key switch points in complex life cycles such as hatching and metamorphosis have stressed the importance of considering multiple stages, most empirical work has focused on a single life stage. However, the relationship between the fitness components of different life stages may be complex. Ontogenetic switch points such as hatching and metamorphosis do not represent new beginnings—carryover effects across stages can arise when environmental effects on the density and/or traits of early ontogenetic stages subsequently alter mortality or growth in later stages. In this study, I examine the effects of egg- and larval-stage predators on larval performance, size at metamorphosis, and post-metamorphic predation in the African tree frog Hyperolius spinigularis. I monitored the density and survival of arboreal H. spinigularis clutches in the field to estimate how much egg-stage predation reduced the input of tadpoles into the pond. I then conducted experiments to determine: (1) how reductions in initial larval density due to egg predators affect larval survival and mass and age at metamorphosis in the presence and absence of aquatic larval predators, dragonfly larvae, and (2) how differences in mass or age at metamorphosis arising from predation in the embryonic and larval environments affect encounters with post-metamorphic predators, fishing spiders. Reduction in larval densities due to egg predation tended to increase per capita larval survival, decrease larval duration and increase mass at metamorphosis. Larval predators decreased larval survival and had density-dependent effects on larval duration and mass at metamorphosis. The combined effects of embryonic and larval-stage predators increased mass at metamorphosis of survivors by 91%. Larger mass at metamorphosis may have immediate fitness benefits, as larger metamorphs had higher survival in encounters with fishing spiders. Thus, the effects of predators early in ontogeny can alter predation risk even two life stages later.     

Density reductions by predatory trout increase adult size and fecundity of surviving caddisfly larvae in a detritus-based stream food web

1. In some situations, individuals surviving in environments where predation is intense can grow faster because the benefits of release from intraspecific competition outweigh costs associated with anti-predator responses. Whether these 'thinning' effects of predation occur in detritus-based food webs where resource renewal occurs independently of consumption by consumers was studied. We investigated how effects of predatory brown trout ( Salmo trutta ) on the larvae of the detritivorous stream caddisfly, Zelandopsyche ingens , influenced the size and fecundity of the caddisfly adults.
2. Trout substantially reduced the abundance of Z. ingens larvae, but adult male and female Z. ingens were significantly larger in trout streams compared to fishless streams. Females in trout streams had 33% more eggs than fishless stream females, and egg sizes were not significantly different. In mesocosms, Z. ingens larvae in low density treatments reflecting trout stream abundances grew significantly faster than larvae in high density treatments that were characteristic of fishless stream abundances. Non-lethal trout presence did not influence case building behaviour, feeding rates or growth or Z. ingens larvae, indicating non-lethal effects of predators were negligible.
3. Increased adult size and fecundity associated with trout stream individuals were probably a result of predator thinning of larval density indirectly releasing surviving Z. ingens from intraspecific competition. Thus, predator thinning did influence interactions between larvae in this detritus-based food web as larval growth was strongly density-dependent. However, extrapolating the total number of eggs potentially produced indicates the increased fecundity of females in trout streams would not compensate for losses of larvae to trout predation.     

Ecological Risk Aversion and Foraging Behaviors of Juvenile Squirrel Monkeys (Saimiri sciureus)

The ‘ecological risk aversion hypothesis’ [C.H. Janson and C.P. van Schaik, Juvenile Primates, Oxford Univ. Press, New York (1993), pp. 57–74] proposes that the pattern of slow growth characteristic of juvenile primates is a response to ecological risks (predation and starvation) experienced by juveniles. Juveniles are thought to avoid predation risk by positioning themselves near conspecifics, therefore experiencing high levels of feeding competition with older individuals, reduced access to resources and, consequently, high starvation risks during periods of food scarcity. The present study compared the foraging behaviors of juvenile and adult squirrel monkeys, a small neotropical primate characterized by a long juvenile period, to determine how predation and starvation risks affected juvenile behaviors. The study was conducted in Eastern Amazonia, in a seasonal environment. Due to their slow development, small body size and large group sizes, it was expected that juveniles in this species would behave in a manner consistent with the risk aversion hypothesis. However, age differences in foraging efficiency and foraging success were smaller than predicted. There was also no evidence that juveniles sacrificed access to food for predator protection. Adults did not have preferential access to fruit patches and direct competition was rare. Feeding competition for prey, the most common resource in the troop's diet, was negligible. Therefore, the slow growth and long juvenile period of squirrel monkeys do not correspond with evidence of predation or starvation risk, as predicted by the risk aversion hypothesis.     

Strong influences of larval diet history on subsequent post-settlement growth in the freshwater mollusc Dreissena polymorpha

A significant seasonal variation in size at settlement has been observed in newly settled larvae of Dreissena polymorpha in Lake Constance. Diet quality, which varies temporally and spatially in freshwater habitats, has been suggested as a significant factor influencing the life history and development of freshwater invertebrates. Accordingly, experiments were conducted with field-collected larvae to test the proposal that diet quality can determine planktonic larval growth rates, size at settlement and subsequent post-metamorphic growth rates. Larvae were fed one of two diets or starved. One diet was composed of cyanobacterial cells, which are deficient in polyunsaturated fatty acids (PUFAs) and the other was a mixed diet rich in PUFAs. Freshly metamorphosed animals from the starvation treatment had a carbon content per individual 70% lower than that of larvae fed the mixed diet. This apparent exhaustion of larval internal reserves resulted in a 50% reduction of the post-metamorphic growth rates. Growth was also reduced in animals previously fed the cyanobacterial diet. Hence, low food quantity or low food quality during the larval stage of D. polymorpha, lead to irreversible effects for post-metamorphic animals and are related to inferior competitive abilities.     

Temporal environmental variation and phenotypic plasticity: a mechanism underlying priority effects

Understanding the role of history in the formation of communities has been a major challenge in community ecology. Here, we explore the role of phenotypic plasticity and its associated trait‐mediated indirect interactions as a mechanism behind priority effects. Using organisms with inducible defenses as a model system, we examine how aquatic communities initially containing different predator environments are affected at the individual and community level by the colonization of a second predator. Snails and tadpoles were established in four different caged‐predator environments (no predator, fish, crayfish or water bugs). These four communities were then crossed with three predator colonization treatments (no colonization, early colonization, or late colonization) using lethal water bugs as the predator. The snails responded to the caged predator environments with predator‐specific behavioral and morphological defenses. In the colonization treatments, snails possessing the wrong phenotype attempted to induce phenotypic changes to defend themselves against the new risk. However, snails initially induced by a different predator environment often suffered high predation rates. Hence, temporal variation in predation risk not only challenged the snail prey to try to track this environmental variation through time by adjusting their defensive phenotypes, but also caused trait‐mediated interactions between snails and the colonizing predator. For tadpoles within these communities, there was little evidence that the morphological responses of snails indirectly effected tadpole predation rates by colonizing water bugs. Unexpectedly, predation rates on tadpoles by colonizing water bugs were generally higher in the three caged‐predator treatments, suggesting that water bugs elevated their foraging activity in response to potentially competing predators. In summary, we demonstrate an important priority effect in which the initial occurrence of one species of predator can facilitate predation by a second predator that colonizes at a later date (i.e. a TMII) suggesting that phenotypic plasticity can be an important driver behind priority effects (i.e. historical exposure to predators).     

Immune deployment increases larval vulnerability to predators and inhibits adult life‐history traits in a dragonfly

While deploying immune defences early in ontogeny can trade‐off with the production and maintenance of other important traits across the entire life cycle, it remains largely unexplored how features of the environment shape the magnitude or presence of these lifetime costs. Greater predation risk during the juvenile stage may particularly influence such costs by (1) magnifying the survival costs that arise from any handicap of juvenile avoidance traits and/or (2) intensifying allocation trade‐offs with important adult traits. Here, we tested for predator‐dependent costs of immune deployment within and across life stages using the dragonfly, Pachydiplax longipennis. We first examined how larval immune deployment affected two traits associated with larval vulnerability to predators: escape distance and foraging under predation risk. Larvae that were induced to mount an immune response had shorter escape distances but lower foraging activity in the presence of predator cues. We also induced immune responses in larvae and reared them through emergence in mesocosms that differed in the presence of large predatory dragonfly larvae (Aeshnidae spp.). Immune‐challenged larvae had later emergence overall and lower survival in pools with predators. Immune‐challenged males were also smaller at emergence and developed less sexually selected melanin wing coloration, but these effects were independent of predator treatment. Overall, these results highlight how mounting an immune defence early in ontogeny can have substantial ecological and physiological costs that manifest both within and across life stages.     

Ontogenetic changes in the association between antipredator responses and growth variables

Abstract.  1. An organism's growth parameters are expected to depend on environmental constraints, such as predation risk and food supply. However, antipredator responses, food intake, and thus growth of an animal may be mediated by behavioural traits, which are likely to differ among developmental stages. In this study, it was investigated how the relationship between growth and behavioural antipredator responses changes during ontogeny in the time-constrained dragonfly species Libellula depressa , and which factors influenced specific behavioural decisions at different points in ontogeny.
2. The results revealed that behavioural strategies differed between larval developmental sages, depending on associations between larval growth, food supply, and predation risk. Early in ontogeny, faster development was correlated with high larval activity and high food supply. This resulted in high activity levels under high food conditions irrespectively of predator presence, and under low food supply in predator absence only. In the intermediate stage of development, all larvae displayed a high activity level, which was correlated in general with fast development. However, growth later in ontogeny was not only influenced by the activity level, but also by predator presence and food supply, with larvae reared under high food supply and/or in presence of predators attaining a higher final mass. Thus, not only the way in which larval growth parameters and behaviour are related changed during development, but also whether the factors influenced larval growth and behaviour. Once the larvae reached the ultimate stage of development, in which they overwinter, behavioural patterns observed were consistent with model predictions.
3. It is advocated that behavioural plasticity of prey organisms in different developmental stages should be analysed in the context of associated growth variables.     

Phenotypic plasticity in complex environments: effects of structural complexity on predator‐ and competitor‐induced phenotypes of tadpoles of the wood frog,Rana sylvatica

Theoretical and empirical research has demonstrated that phenotypically plastic responses to one environment are dependent on other environmental attributes. Such research is critical considering the complexity of natural habitats, yet few studies have examined how multiple environments affect patterns of plasticity and the adaptiveness of the resulting phenotypes within complex habitats. The present study examines how wood frog (Rana sylvatica) tadpoles alter their behavioural and morphological phenotypes in response to predation risk from larval diving beetles (Dytiscus spp.), competition from conspecifics, and physical structural complexity. It also tests whether structure affects selection intensities by Dytiscus larvae on tadpole morphological traits. Predation risk and competition induced typical changes to tadpole behaviour and morphology. Structure did not induce changes to any phenotype, nor did it interact with predation risk or competition in affecting phenotypes. Furthermore, structure did not affect the predator selection intensities on any morphological trait. Dytiscus larvae selected for shallow, short tailfins, and large tail muscles, yet tadpoles only developed deep tail muscles when raised in the presence of predator cues. These apparently maladaptive responses may have been a result of correlations between phenotypes. The present study expands plasticity research by examining the adaptiveness of plastic responses in complex environments. Additionally, the present study demonstrates that not all environments induce plastic responses. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 853–863.     

Effects of a larval antipredator response and larval diet on adult phenotype in an aposematic ladybird beetle

Many ladybird beetles respond to a potential predation event by `reflex bleeding' or secreting a noxious defensive chemical that is similar to hemolymph. Both adults and larvae show this response. Reflex bleeding is known to reduce predator attack rates and increase prey survival after an attack, especially when reflex bleeding is employed in combination with other cues such as odor and warning coloration. In this experiment, we examined how variability in the number of reflex bleeding events and food quality in the larval stage of the aposematic ladybird beetle Harmonia axyridis affected elytral color, development time, and terminal size in adults. Effects of reflex bleeding were subtle and may have been influenced by diet treatments. Adult color did not differ between bleed treatment groups but beetles that reflex bled tended to take longer to develop and grow to smaller sizes than control group beetles. There were clear and strong effects of larval diet on adult phenotype: an ad libitum pollen diet resulted in paler adult coloration, shorter development time, and larger adult size relative to a limited-availability aphid diet. Our results suggest that the best environment for producing bright-red coloration may not be the best environment for favorable expression of life history characters, especially under stressful conditions. Interactions between different life history stages of H. axyridis are also discussed. Received: 20 April 1997 / Accepted: 30 September 1997