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The optimal timing for initiating reproduction (i.e., the age at first reproduction) is a critical life history trait describing aspects of an individual's resource-allocation strategy. Recent theoretical and empirical work has demonstrated that this trait is also tied to mating system expression when individuals have the opportunity to reproduce via both self-fertilization and cross-fertilization. A strategy of "delayed selfing" has emerged as a "best of both worlds" arrangement where, in the absence of a mate, an individual will delay reproduction (selfing) to "wait" for a mate. Herein, we extend previously developed predictive optimization models for the timing of reproduction to a situation where organisms can allocate their resources to size-dependent and size-independent defensive strategies to counter the threat of predation. By incorporating inducible defenses into a predictive framework for analyzing life history expression and evolution, we can more accurately evaluate the role that allocation strategy plays in altering the optimal waiting time. We compare our model to previous models and empirical results highlighting that incorporation of inducible defenses into the model broadens the parameter space in which a waiting time is expected and often leads to a predicted waiting time that is longer than in the situation without inducible defenses. In particular, a waiting time is predicted to exist regardless of the strength of inbreeding depression in the population.

Because establishing a new population often depends critically on finding mates, individuals capable of uniparental reproduction may have a colonization advantage. Accordingly, there should be an over-representation of colonizing species in which individuals can reproduce without a mate, particularly in isolated locales such as oceanic islands. Despite the intuitive appeal of this colonization filter hypothesis (known as Baker's law), more than six decades of analyses have yielded mixed findings. We assembled a dataset of island and mainland plant breeding systems, focusing on the presence or absence of self-incompatibility. Because this trait enforces outcrossing and is unlikely to re-evolve on short timescales if it is lost, breeding system is especially likely to reflect the colonization filter. We found significantly more self-compatible species on islands than mainlands across a sample of > 1500 species from three widely distributed flowering plant families (Asteraceae, Brassicaceae and Solanaceae). Overall, 66% of island species were self-compatible, compared with 41% of mainland species. Our results demonstrate that the presence or absence of self-incompatibility has strong explanatory power for plant geographical patterns. Island floras around the world thus reflect the role of a key reproductive trait in filtering potential colonizing species in these three plant families.

Senescence is not a static property of an individual or population, but rather it is a dynamic process that may be influenced by environmental conditions. This can occur in at least two ways: in the long-term across multiple generations, and in the short-term via phenotypic plasticity. The former has attracted a lot of attention, both theoretically and empirically; the latter has lagged behind. To determine whether two important environmental variables (predation risk and mate availability) affect the pattern of actuarial senescence (i.e., the increase in mortality with age), we reared 30 full-sib families of the simultaneously hermaphroditic freshwater snail Physa acuta under four different environmental conditions and tracked individuals until death. Individuals were reared in a 2x2 factorial experiment that manipulated the non-lethal presence of chemical cues from predatory crayfish (presence/absence) and the opportunity to mate with an unrelated partner (mated/not mated). Snails that receive a partner reproduce by outcrossing, while those that remain in isolation can reproduce by self-fertilization. We compared the cumulative survival curves to test for an effect of predation risk and mating. The hazard ratio for the predation-risk comparison was 1.042 indicating no significant difference between the curves. However, the hazard ratio for the mating comparison was 4.021, reflecting a significant reduction in survival probability for mated snails relative to isolated snails. As such, mating resulted in a much shorter lifespan, an outcome that we interpret in terms of shifting resource allocation. This article is protected by copyright. All rights reserved.

Reproductive performance is often age-dependent, showing patterns of improvement and/or senescence as well as trade-offs with other traits throughout the lifespan. High levels of extrinsic mortality (e.g., from predators) have been shown to sometimes, but not always, select for accelerated actuarial senescence in nature and in the lab. Here, we explore the inductive (i.e., plastic) effects of predation risk (i.e., nonlethal exposure to chemical cues from predators) on the reproductive success of freshwater snails (Physa acuta). Snails were reared either in the presence or absence of chemical cues from predatory crayfish and mated early in life or late in life (a 2 × 2 factorial design); we measured egg hatching and early post-hatching survival of their offspring. Both age and predation risk reduced reproductive success, illustrating that predation risk can have a cross-generational effect on the early survival of juveniles. Further, the decline in reproductive success was over three times faster under predation risk compared to the no-predator treatment, an effect that stemmed from a disproportionate, negative effect of predation risk on the post-hatching survival instead of hatching rate. We discuss our results in terms of a hypothesized consequence of elevated stress hormone levels.

Reproductive success is a critical fitness attribute that is directly influenced by resource availability. Here, we investigate the effects of diet-based resource availability on three interrelated aspects of reproductive success: a change in mating system based on mate availability, consequent inbreeding depression, and the deterioration of reproductive efficiency with age (senescence). We employed a factorial experimental design using 22 full-sib families of the hermaphroditic freshwater snail Physa acuta to explore these interactions. Individual snails were reared in one of two mate-availability treatments (isolated [selfing] or occasionally paired [outcrossing]) and one of two diet treatments (boiled lettuce or Spirulina, an algae that is rich in protein, vitamins, and minerals). Spirulina-fed snails initiated reproduction at a 13% earlier age and 7% larger size than lettuce-fed snails. Spirulina also resulted in a 30% reduction in the time delay before selfing. Compared to lettuce, a diet of Spirulina increased inbreeding depression by 52% for egg hatching rate and 64% for posthatching juvenile survival. Furthermore, Spirulina led to a 15-fold increase in the rate of reproductive senescence compared with a diet of lettuce. These transgenerational, interactive effects of diet on inbreeding depression and reproductive senescence are discussed in the context of diet-induced phenotypic plasticity.

Predation risk is known to induce numerous, often adaptive, plastic changes in the morphology, life history, and behavior of prey. However, little is known about how abiotic factors such as environmental nutrient availability affect these responses. In this experiment, a freshwater pulmonate snail (Physa acuta) was used to determine how predator-induced shell morphology was affected by the availability of an abiotic nutrient important for shell growth (calcium). Each snail was reared individually in a factorial experiment consisting of two predator treatments (chemical cues from crayfish present or absent) and three calcium concentrations (20, 45, or 70 mg/L). Calcium induced larger, heavier, and thicker shells, while predator cues induced heavier and thicker shells. Importantly, the magnitude of the predator effect on shell thickness (i.e., the inducible defense) increased with additional calcium availability. Our results demonstrate the importance of nutrient availability in mediating and limiting the amount of adaptive predator-induced plasticity.

Summary Traditionally, many breeding traits (e.g. the timing and size of clutches) were considered to be female-only traits in that males played little-to-no role in their expression. Although the contribution of males to such breeding traits, as well as other aspects of reproduction, is increasingly recognized, few studies have demonstrated the effects of male age and life history on breeding traits and, importantly, whether these effects are underlined by additive-genetic variation. Here, we take advantage of a long-term data set on mute swans (Cygnus olor) to demonstrate that the ages of both the male and female parents play significant roles in the timing and size of clutches, although recruitment success did not show similar effects. Individual males varied significantly in their influence on the timing of egg laying. We decomposed this variation using an ‘animal model’; competing models that were the source of this variation as additive-genetic or permanent-environmental variation was not statistically distinguishable. Our results add to the growing evidence that reproductive performance should be considered as a product of the identity and condition of both parents.

Snails (Mollusca: Gastropoda) are a diverse class of mollusks inhabiting marine, terrestrial, and freshwater ecosystems. Although the most common reproductive system is gonochorism, an astounding diversity of other systems have repeatedly evolved, including apomictic parthenogenesis, protandrous hermaphroditism, and simultaneous hermaphroditism. Some snails have evolved specific mechanisms of sex determination (e.g., sex chromosomes), while others show that environmental contaminants can induce changes in reproductive anatomy (e.g., imposex). Despite the diversity of reproductive systems and strategies that has been observed, and the fact that Gastropods serve as model systems for addressing a wide range of questions, a great deal remains to be discovered.

The existence of an evolutionary syndrome linking dispersal and mating-system traits has been discussed in both plants and animals. In animals, dispersal as a means of inbreeding-avoidance has been cited as an ultimate cause of sex-biased dispersal. In plants, self-compatibility is widespread, which is often cited as a mechanism for reproductive assurance in organisms that have limited control of dispersal. Limited dispersal has also been hypothesized to minimize outbreeding depression and increase local adaptation. Here, we compare and contrast the various evolutionary hypotheses that link dispersal and the mating system in both plants and animals. We conclude that the majority of theoretical evidence supports the existence of two evolutionary syndromes: (1) outcrossing and dispersing; (2) inbreeding and not dispersing. In the light of the evidence compiled, we advocate for a redefinition of Baker’s law, which we consider to be an exception rather than the rule. As environmental heterogeneity is common in nature, the role of bet-hedging in the evolution of these strategies is likely to be very strong, albeit difficult to prove empirically. Lastly, we argue that exceptions to these two general syndromes (e.g., inbreeding and dispersing) should not be viewed as contradictory; instead they merit exceptional attention because they represent unusual biological adaptations. Throughout, we refer to specific empirical examples to illustrate these scenarios and conclude by suggesting that a meta-analysis of the available data would be a useful next step.

In hermaphrodites, traits that influence the selfing rate can coevolve with inbreeding depression, leading to the emergence of evolutionary syndromes. Theory predicts a negative correlation between inbreeding depression and selfing rate across species. This prediction has only been examined and validated in vascular plants. Furthermore, selfing rates are often influenced by environmental conditions (e.g., lack of mates or pollinators), and species are predicted to evolve mechanisms to buffer this variation. We extend previous studies of mating-system syndromes in two ways. First, we assembled a new dataset on Basommatophoran snails (17 species, including new data on 12 species). Second, we measured how species responded to variation in mate availability. Specifically, we quantified the waiting time before selfing (i.e., how long the onset of reproduction is delayed in the absence of mates). Selfing rates were negatively correlated with both inbreeding depression and the waiting time. Species with stronger inbreeding depression exhibited longer waiting times. These patterns obtained on Basommatophorans still hold when including eight other hermaphroditic animals. Our results support the hypothesis that selection drives the evolution of mating-system syndromes in animals. The reaction norm of selfing rates to mate availability is a key target of natural selection in this context. © 2011 The Author(s). Evolution© 2011 The Society for the Study of Evolution.

Environmental effects on mating system expression are central to understanding mating system evolution in nature. Here, I report the results from a quantitative-genetic experiment aimed at understanding the role of predation risk in the expression and evolution of life-history and mating-system traits in a hermaphroditic freshwater snail (Physa acuta). I reared 30 full-sib families in four environments that factorially contrast predation risk and mate availability and measured age/size at first reproduction, growth rate, a morphological defense, and the early survival of outcrossed/selfed eggs that were laid under predator/no-predator conditions. I evaluated the genetic basis of trade-offs among traits and the stability of the G matrix across environments. Mating reduced growth while predation risk increased growth, but the effects of mating were weaker for predator-induced snails and the effects of predation risk were weaker for snails without mates. Predation risk reduced the amount of time that individuals waited before self-fertilizing and reduced inbreeding depression in the offspring. There was a positive among-family relationship between the amount of time that individuals delayed selfing under predation risk and the magnitude of inbreeding depression. These results highlight several potential roles of enemies in mating-system expression and evolution.

Environmental effects on the evolution of mating systems are increasingly discussed, but we lack many examples of how environmental conditions affect the expression and consequences of alternative mating systems. Variation in mate availability sets up a trade-off between reproductive assurance and inbreeding depression, but the consequences of both mate limitation and inbreeding may depend on other environmental conditions. Predation risk is common under natural conditions, and known to affect allocation to reproduction, but we know little about the effects of isolation and inbreeding under predation risk. We reared selfed and outcrossed hermaphroditic freshwater snails (Physa acuta) in four environments (predator cues present or absent crossed with mating partners available or not) and quantified life-history traits and cumulative lifetime fitness. Our results confirm that isolation from mates can increase longevity and growth, resulting in higher lifetime fecundity. Thus, we observed no evidence for mate limitation of reproduction. However, reproduction under isolation (i.e., selfing) resulted in inbreeding depression, which should counteract the benefits of selfing. Inbreeding depression in fitness occurred in both predator and no-predator environments, but there was no overall change in inbreeding depression with predator cues. This represents, to our knowledge, the first empirical estimate of the effect of predation risk on inbreeding depression in an animal. Cumulative fitness was most influenced by early survival and especially early fecundity. As predation risk and inbreeding (both ancestral and due to a lack of mates) reduced early fecundity, these effect are predicted to have important contributions to population growth under natural conditions. Therefore life-history plasticity (e.g., delayed reproduction) is likely to be very important to overall fitness.

When the optimal phenotype differs among environments, adaptive phenotypic plasticity can evolve unless constraints impede such evolution. Costs and limits of plasticity have been proposed as important constraints on the evolution of plasticity, yet confusion exists over their distinction. We attempt to clarify these concepts by reviewing their categorization and measurement, highlighting how costs and limits are defined in different currencies (and may describe the same phenomenon). Conclusions from studies that measure the costs of plasticity have been equivocal, but we caution that these conclusions may be premature owing to a potentially common correlation between environment-specific trait values and the magnitude of trait plasticities (i.e. multi-collinearity) that results in imprecise and/or biased estimates of the costs. Meanwhile, our understanding of the limits of plasticity, and how they may be underlain by the costs of plasticity, is still in its infancy. Based on our re-evaluation of these constraints, we discuss areas for future research.

Although a large portion of plant and animal species exhibit intermediate levels of outcrossing, the factors that maintain this wealth of variation are not well understood. Natural enemies are one relatively understudied ecological factor that may influence the evolutionary stability of mixed mating. In this paper, we aim for a conceptual unification of the role of enemies in mating system expression and evolution in both hermaphroditic animals and plants. We review current theory and detail the potential effects of enemies on fundamental mating system parameters. In doing so, we identify situations in which consideration of enemies alters expectations about the stability of mixed mating. Generally, we find that inclusion of the enemy dimension may broaden conditions in which mixed mating systems are evolutionarily stable. Finally, we highlight avenues ripe for future theoretical and empirical work that will advance our understanding of enemies in the expression and evolution of mixed mating in their hosts/victims, including examination of feedback cycles between victims and enemies and quantification of mating system-related parameters in victim populations in the presence and absence of enemies.

Baker's law refers to the tendency for species that establish on islands by long-distance dispersal to show an increased capacity for self-fertilization because of the advantage of self-compatibility when colonizing new habitat. Despite its intuitive appeal and broad empirical support, it has received substantial criticism over the years since it was proclaimed in the 1950s, not least because it seemed to be contradicted by the high frequency of dioecy on islands. Recent theoretical work has again questioned the generality and scope of Baker's law. Here, we attempt to discern where the idea is useful to apply and where it is not. We conclude that several of the perceived problems with Baker's law fall away when a narrower perspective is adopted on how it should be circumscribed. We emphasize that Baker's law should be read in terms of an enrichment of a capacity for uniparental reproduction in colonizing situations, rather than of high selfing rates. We suggest that Baker's law might be tested in four different contexts, which set the breadth of its scope: the colonization of oceanic islands, metapopulation dynamics with recurrent colonization, range expansions with recurrent colonization, and colonization through species invasions.

Phenotypic plasticity is ubiquitous and generally regarded as a key mechanism for enabling organisms to survive in the face of environmental change. Because no organism is infinitely or ideally plastic, theory suggests that there must be limits (for example, the lack of ability to produce an optimal trait) to the evolution of phenotypic plasticity, or that plasticity may have inherent significant costs. Yet numerous experimental studies have not detected widespread costs. Explicitly differentiating plasticity costs from phenotype costs, we re-evaluate fundamental questions of the limits to the evolution of plasticity and of generalists vs specialists. We advocate for the view that relaxed selection and variable selection intensities are likely more important constraints to the evolution of plasticity than the costs of plasticity. Some forms of plasticity, such as learning, may be inherently costly. In addition, we examine opportunities to offset costs of phenotypes through ontogeny, amelioration of phenotypic costs across environments, and the condition-dependent hypothesis. We propose avenues of further inquiry in the limits of plasticity using new and classic methods of ecological parameterization, phylogenetics and omics in the context of answering questions on the constraints of plasticity. Given plasticity's key role in coping with environmental change, approaches spanning the spectrum from applied to basic will greatly enrich our understanding of the evolution of plasticity and resolve our understanding of limits.

The laminar scales of the fern Elaphoglossum paleaceum were studied to determine if they act as a morphological mechanism to protect leaves from excess light. We hypothesized that if scales serve this purpose, then individual plants growing in high light would have greater laminar scale density than those in low light. For our first experiment, plants from high light roadsides were collected and subjected to artificial scale removal and then exposed to super-saturating pulse of light for 30 min. We found that leaves with their scales removed exhibited significantly greater photoinhibition than leaves with scales intact. Leaves with intact scales recovered fully after twelve hours whereas those with scales removed remained photoinhibited. Scale density on the adaxial side of leaves was positively correlated with light intensity. The data from this study indicate that fern laminar scales help reduce photoinhibition and potentially function as a morphological defense against photodamage.

Studies of phenotypic plasticity frequently demonstrate functional trade-offs between alternative phenotypes by documenting environment-specific costs and benefits. However, the functional mechanisms underlying these trade-offs are often unknown. For example, predator-induced traits typically provide superior predator resistance but slower growth, while competitor-induced traits provide better growth but inferior predator resistance. While the mechanisms underlying predator resistance have been identified, the mechanisms underlying differential growth have remained elusive. To determine whether competitor and predator environments affect individual growth by induced changes in foraging morphology, we raised wood frog tadpoles (Rana sylvatica) under a factorial combination of competitors and predators and assessed changes in mouthparts that might affect growth. In general, competitors induced relatively larger oral discs, wider beaks, and longer tooth rows, while predators induced relatively smaller oral discs, narrower beaks, and shorter tooth rows. These effects were interactive; the largest competitor-induced responses occurred under high predator density and the largest predator-induced responses occurred under low competition. Further, one of the tooth rows that commonly appeared under low predation risk was frequently absent under high predation risk. These discoveries suggest that predator and competitor environments can have profound effects on prey foraging structures and that these effects set up growth trade-offs between phenotypes that favor the evolution of phenotypically plastic responses.

Predators commonly induce phenotypic changes that make prey better at surviving predation at the cost of reduced growth. While we have a good understanding of how trait changes affect predation risk, we lack a mechanistic understanding of why predator-induced phenotypes differ in growth. Using two mesocosm experiments, we combined phenotypic plasticity theory with predictions from optimal digestion theory to demonstrate that intra- and interspecific competition induced relatively long guts while predators induced relatively short guts. The longer guts induced by competition appear to be an adaptive response that allows more efficient digestion and more rapid growth whereas the shorter guts induced by predators appear to result from a tradeoff of building larger tails in predator environments at the cost of smaller bodies. By combining these two bodies of theory, we now have a much better understanding of the mechanisms that cause the phenotypic trade-offs that select for inducible defences.

The diversity of reproductive strategies in nature is shaped by a plethora of factors including energy availability. For example, both low temperatures and limited food availability could increase larval exposure to predation by slowing development, selecting against pelagic and/or feeding larvae. The frequency of hermaphroditism could increase under low food availability as population density (and hence mate availability) decreases. We examine the relationship between reproductive/life-history traits and energy availability for 189 marine gastropod families. Only larval type was related to energy availability with the odds of having planktotrophic larvae versus direct development decreasing by 1% with every one-unit increase in the square root of carbon flux. Simultaneous hermaphroditism also potentially increases with carbon flux, but this effect disappears when accounting for evolutionary relationships among taxa. Our findings are in contrast to some theory and empirical work demonstrating that hermaphroditism should increase and planktotrophic development should decrease with decreasing productivity. Instead, they suggest that some reproductive strategies are too energetically expensive at low food availabilities, or arise only when energy is available, and others serve to capitalize on opportunities for aggregation or increased energy availability.

It is commonly observed that reproduction decreases with age, often at a different rate in males and females. This phenomenon is generally interpreted as senescence. Such reproductive declines may stem from at least two sources: a change in resource allocation and a decline in the ability to convert resources into offspring. This distinction is important because a shift in resource allocation may be favoured by selection, while reduced efficiency is purely deleterious. We propose a way to distinguish whether a decline in reproduction is purely deleterious based on estimating reproductive investment, output, and their ratio, efficiency. We apply this approach to the hermaphroditic snail Physa acuta and demonstrate that both male and female functions decline with age. The male decline largely stems from reduced investment into male activity while female decline is due to increased reproductive inefficiency. This shows that age-related declines in reproduction can occur for a number of different reasons, a distinction that is usually masked by the general term ‘senescence’. This approach could be applied to any species to evaluate age-related reproductive decline. We advocate that future studies measure age trajectories of reproductive investment and output to explore the potential processes hidden behind the observation that reproduction declines with age.

Summary Seed dormancy, by controlling the timing of germination, can strongly affect plant survival. The kind of seed dormancy, therefore, can influence both population and species-level processes such as colonization, adaptation, speciation, and extinction. We used a dataset comprising over 14 000 taxa in 318 families across the seed plants to test hypotheses on the evolution of different kinds of seed dormancy and their association with lineage diversification. We found morphophysiological dormancy to be the most likely ancestral state of seed plants, suggesting that physiologically regulated dormancy in response to environmental cues was present at the origin of seed plants. Additionally, we found that physiological dormancy (PD), once disassociated from morphological dormancy, acted as an ‘evolutionary hub’ from which other dormancy classes evolved, and that it was associated with higher rates of lineage diversification via higher speciation rates. The environmental sensitivity provided by dormancy in general, and by PD in particular, appears to be a key trait in the diversification of seed plants.

Understanding the evolution of reaction norms remains a major challenge in ecology and evolution. Investigating evolutionary divergence in reaction norm shapes between populations and closely related species is one approach to providing insights. Here we use a meta-analytic approach to compare divergence in reaction norms of closely related species or populations of animals and plants across types of traits and environments. We quantified mean-standardized differences in overall trait means (Offset) and reaction norm shape (including both Slope and Curvature). These analyses revealed that differences in shape (Slope and Curvature together) were generally greater than differences in Offset. Additionally, differences in Curvature were generally greater than differences in Slope. The type of taxon contrast (species vs. population), trait, organism, and the type and novelty of environments all contributed to the best-fitting models, especially for Offset, Curvature, and the total differences (Total) between reaction norms. Congeneric species had greater differences in reaction norms than populations, and novel environmental conditions increased the differences in reaction norms between populations or species. These results show that evolutionary divergence of curvature is common and should be considered an important aspect of plasticity, together with slope. Biological details about traits and environments, including cryptic variation expressed in novel environmental conditions, may be critical to understanding how reaction norms evolve in novel and rapidly changing environments.

There are many different ways to measure phenotypic plasticity [[1][1]]. In the endeavour to measure the costs of plasticity, researchers often employ large quantitative-genetic experimental designs that rear related groups of organisms in two environments. Under such two-environment designs

Sexual selection is often quantified using Bateman gradients, which represent sex-specific regression slopes of reproductive success on mating success and thus describe the expected fitness returns from mating more often. Although the analytical framework for Bateman gradients aimed at covering all sexual systems, empirical studies are biased toward separate-sex organisms, probably because important characteristics of other systems remain incompletely treated. Our synthesis complements the existing Bateman gradient approach with three essential reproductive features of simultaneous hermaphrodites. First, mating in one sex may affect fitness via the opposite sex, for example, through energetic trade-offs. We integrate cross-sex selection effects and show how they help characterizing sexually mutualistic versus antagonistic selection. Second, male and female mating successes may be correlated, complicating the interpretation of Bateman gradients. We show how to quantify the impact of this correlation on sexual selection and propose a principal component analysis on male and female mating success to facilitate interpretation. Third, self-fertilization is accounted for by adding selfed progeny as a separate category of reproductive success to analyses of Bateman gradients. Finally, using a worked example from the snail Biomphalaria glabrata, we illustrate how the extended analytical framework can enhance our understanding of sexual selection in hermaphroditic animals and plants. © 2010 by The University of Chicago.

While much attention has been paid to the effects of inbreeding on fitness, this has mostly come from a genetic perspective. Consequently, the interaction between inbreeding and the environment is less well understood. To understand the effects of inbreeding in natural populations where environmental conditions are variable, we need to examine not only how the effects of inbreeding change among environments but also how inbreeding may affect the ability to respond to environmental conditions (i.e. phenotypic plasticity). We reared selfed and outcrossed hermaphroditic snails (Physa acuta) in the presence and absence of chemical cues from predatory crayfish and quantified expression of an inducible defence, an adaptively plastic response to predation risk. Overall, inbred snails exhibited reduced defences, but more importantly, inbreeding reduced the expression of predator-induced adaptive plasticity. Inbreeding depression in defensive morphology was 26 per cent and inbreeding depression in the plasticity of this trait was 48 per cent. Inbreeding depression in adaptive plasticity may be important to understanding the effects of inbreeding in nature.

Encountering mates and avoiding predators are ubiquitous challenges faced by many organisms and they can affect the expression of many traits including growth, timing of maturity and resource allocation to reproduction. However, these two factors are commonly considered in isolation rather than simultaneously. We examined whether predation risk and mate availability interact to affect morphology and life-history traits (including lifetime fecundity) of a hermaphroditic snail (Physa acuta). We found that mate availability reduced juvenile growth rate and final size. Predator cues from crayfish induced delayed reproduction, but there were no reduced fecundity costs associated with predator induction. Although there were interactive effects on longevity, lifetime fecundity was determined by the number of reproductive days. Therefore, our results indicate a resource-allocation trade-off among growth, longevity and reproduction. Future consideration of this interaction will be important for understanding how resource-allocation plasticity affects the integration of defensive, life-history and mating-system traits.

The last decade has seen an explosion in the number of studies exploring predator-induced plasticity. Recently, there has been a call for more comprehensive approaches that can identify functional relationships between traits, constraints on phenotypic responses, and the cost and benefits of alternative phenotypes. In this study, we exposed Helisoma trivolvis, a freshwater snail, to a factorial combination of three resource levels and five predator environments (no predator, one or two water bugs, and one or two crayfish) and examined ten traits including behavior, morphology, and life history. Each predator induced a unique suite of behavioral and morphological responses. Snails increased near-surface habitat use with crayfish but not with water bugs. Further, crayfish induced narrow and high shells whereas water bugs induced wide shells and wide apertures. In terms of life history, both predators induced delayed reproduction and greater mass at reproduction. However, crayfish induced a greater delay in reproduction that resulted in reduced fecundity whereas water bugs did not induce differences in fecundity. Resource levels impacted the morphology of H. trivolvis; snails reared with greater resource levels produced higher shells, narrower shells, and wider apertures. Resource levels also impacted snail life history; lower resources caused longer times to reproduction and reduced fecundity. Based on an analysis of phenotypic correlations, the morphological responses to each predator most likely represent phenotypic trade-offs. Snails could either produce invasion-resistant shells for defense against water bugs or crush-resistant shells for defense against crayfish, but not both. Our use of a comprehensive approach to examine the responses of H. trivolvis has provided important information regarding the complexity of phenotypic responses to different environments, the patterns of phenotypic integration across environments, and the potential costs and benefits associated with plastic traits.

Excluding insects, hermaphroditism occurs in about one-third of animal species, providing numerous opportunities for the evolution of selfing. Here we provide an overview of reproductive traits in hermaphroditic animal species, review the distribution of selfing rates in animals, and test for ecological correlates of selfing. Our dataset (1342 selfing-rate estimates for 142 species) is 97% based on estimates derived from the analysis of population structure (F(IS)-estimates) using genetic markers. The distribution of selfing is slightly U-shaped and differs significantly from the more strongly U-shaped plant distribution with 47% of animal t-estimates being intermediate (falling between 0.2 and 0.8) compared to 42% for plants. The influence of several factors on the distribution of selfing rates was explored (e.g., number of populations studied per species, habitat, coloniality, sessility, or fertilization type), none of which significantly affect the distribution. Our results suggest that genetic forces might contribute to the evolution of self-fertilization to the same extent in animals and plants, although the high proportion of intermediate outcrossing suggests a significant role of ecological factors (e.g., reproductive assurance) in animals. However, we caution that the distribution of selfing rates in animals is affected by various factors that might bias F(IS)-estimates, including phylogenetic underrepresentation of highly selfing and outcrossing species, various genotyping errors (e.g., null alleles) and inbreeding depression. This highlights the necessity of obtaining better estimates of selfing for hermaphroditic animals, such as genotyping progeny arrays, as in plants.