Life-history plasticity and inbreeding depression under mate limitation and predation risk: cumulative lifetime fitness dissected with a life table response experiment

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.     

鲤幼鱼标准代谢率的个体差异与运动性能和摄食代谢的关系

为考察鲤科鱼类种内个体标准代谢率的差异及其与运动性能和摄食性能的内在关联,本研究以我国广泛分布的鲤(Cyprinids cardio)幼鱼[体重(4.79±0.08)g,n=36]为实验对象,在(25.0±1.0)℃下分别测量实验鱼的标准代谢率(SMR),随后测定单尾鱼的特殊动力作用(SDA)、自发运动、临界游泳速度以及活跃代谢率(MO2active)。实验鱼标准代谢率(SMR)的变幅为76.7~317.6 mg/(kg·h),其变异系数(CV)达24.4%;实验鱼在10 min内的尾鳍摆动次数(P0.05)和摄食代谢峰值(P0.05)均与标准代谢率(SMR)呈正相关;活跃代谢率(MO2active)(P0.05)与摄食代谢峰值以及活跃代谢范围与摄食代谢范围(P0.05)均呈正相关。然而,鲤幼鱼的标准代谢率(SMR)与相对临界游泳速度、活跃代谢率(MO2active)、特殊动力作用(SDA)时间和特殊动力作用(SDA)总量均不相关(所有P0.05)。研究表明,较高标准代谢率(SMR)的鲤幼鱼个体表现较高的活跃性和较强的摄食代谢能力,可能有助于其更易发现食物、逃避天敌以及加快食物处理。     

Feeding while evading predators by a lotic mayfly: linking short-term foraging behaviours to long-term fitness consequences

Laboratory experiments were performed to quantify the combined effects of food abundance (low, high) and predation threat, imposed using a model fish (safe, risky) on fitness correlates (i.e. growth, time to emergence, adult body mass, fecundity, egg size) of the mayfly Baetis tricaudatus. These effects were determined by rearing larvae under different combinations of food abundance and predator threat. Fitness correlates were significantly affected by food abundance, predation threat or the interaction of these factors. High food abundance and the absence of predation threat significantly increased larval growth rates, adult body mass, fecundity, egg size, and decreased time to emergence. Long-term effects of predator threat and food abundance on fitness correlates of B. tricaudatus were compared to previously conducted short-term patch choice trials to test for concordance between short-term measures of patch choice and their potential longer-term fitness consequences. This comparison indicated that patches that were utilized the most were those that yielded the highest fitness benefits. These results suggest that behavioural costs of balancing mortality risk due to predation against food acquisition by B. tricaudatus can have strong fitness consequences.     

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.     

Costs of inducible defence along a resource gradient

In addition to having constitutive defence traits, many organisms also respond to predation by phenotypic plasticity. In order for plasticity to be adaptive, induced defences should incur a benefit to the organism in, for example, decreased risk of predation. However, the production of defence traits may include costs in fitness components such as growth, time to reproduction, or fecundity. To test the hypothesis that the expression of phenotypic plasticity incurs costs, we performed a common garden experiment with a freshwater snail, Radix balthica, a species known to change morphology in the presence of molluscivorous fish. We measured a number of predator-induced morphological and behavioural defence traits in snails that we reared in the presence or absence of chemical cues from fish. Further, we quantified the costs of plasticity in fitness characters related to fecundity and growth. Since plastic responses may be inhibited under limited resource conditions, we reared snails in different densities and thereby levels of competition. Snails exposed to predator cues grew rounder and thicker shells, traits confirmed to be adaptive in environments with fish. Defence traits were consistently expressed independent of density, suggesting strong selection from predatory molluscivorous fish. However, the expression of defence traits resulted in reduced growth rate and fecundity, particularly with limited resources. Our results suggest full defence in predator related traits regardless of resource availability, and costs of defence consequently paid in traits related to fitness.     

Maternal exposure to predation risk decreases offspring antipredator behaviour and survival in threespined stickleback

1. Adaptive maternal programming occurs when mothers alter their offspring's phenotype in response to environmental information such that it improves offspring fitness. When a mother's environment is predictive of the conditions her offspring are likely to encounter, such transgenerational plasticity enables offspring to be better-prepared for this particular environment. However, maternal effects can also have deleterious effects on fitness.2. Here, we test whether female threespined stickleback fish exposed to predation risk adaptively prepare their offspring to cope with predators. We either exposed gravid females to a model predator or not, and compared their offspring's antipredator behaviour and survival when alone with a live predator. Importantly, we measured offspring behaviour and survival in the face of the same type of predator that threatened their mothers (Northern pike).3. We did not find evidence for adaptive maternal programming; offspring of predator-exposed mothers were less likely to orient to the predator than offspring from unexposed mothers. In our predation assay, orienting to the predator was an effective antipredator behaviour and those that oriented, survived for longer.4. In addition, offspring from predator-exposed mothers were caught more quickly by the predator on average than offspring from unexposed mothers. The difference in antipredator behaviour between the maternal predator-exposure treatments offers a potential behavioural mechanism contributing to the difference in survival between maternal treatments.5. However, the strength and direction of the maternal effect on offspring survival depended on offspring size. Specifically, the larger the offspring from predator-exposed mothers, the more vulnerable they were to predation compared to offspring from unexposed mothers.6. Our results suggest that the predation risk perceived by mothers can have long-term behavioural and fitness consequences for offspring in response to the same predator. These stress-mediated maternal effects can have nonadaptive consequences for offspring when they find themselves alone with a predator. In addition, complex interactions between such maternal effects and offspring traits such as size can influence our conclusions about the adaptive nature of maternal effects.     

Predation selects for low resting metabolic rate and consistent individual differences in anti-predator behavior in a beetle

Significant between-individual variation in resting metabolic rate (RMR) of animals is a widespread phenomenon that may have important implications for our understanding of variation in behavior and animal personality. By using wild caught mealworm beetles, Tenebrio molitor, we examined the relationships among survival rate under predator tests, individual response latency time to become immobile under the risk of predation, duration of immobility time, and RMR. Individuals with higher levels of RMR were bold, and bold individuals were found to be more exposed to the risk of bird predation. We found that RMR was positively correlated with the latency of immobility response and negatively correlated with the total duration of immobility. The correlation between behavioral responses suggests a behavioral syndrome in the anti-predator behavior of T. molitor. The results indicate that energy metabolism may be part of a syndrome that involves behavior and life history traits in animals.     

Escape behaviour and ultimate causes of specific induced defences in an anuran tadpole

Induced defences, such as the predator avoidance morphologies in amphibians, result from spatial or temporal variability in predation risk. One important component of this variability should be the difference in hunting strategies between predators. However, little is known about how specific and effective induced defences are to different types of predators. We analysed the impact of both pursuing (fish, Gasterosteus aculeatus) and sit-and-wait (dragonfly, Aeshna cyanea) predators on tadpole (Rana dalmatina) morphology and performance (viz locomotive performance and growth rate). We also investigated the potential benefits of the predator-induced phenotype in the presence of fish predators. Both predators induced deeper tail fins in tadpoles exposed to threat of predation, and stickleback presence also induced longer tails and deeper tail muscles. Morphological and behavioural differences resulted in better escape ability of stickleback-induced tadpoles, leading to improved survival in the face of stickleback predation. These results clearly indicate that specific morphological responses to different types of predators have evolved in R. dalmatina. The specific morphologies suggest low correlations between the traits involved in the defence. Independence of traits allows prey species to fine-tune their response according to current predation risk, so that the benefit of the defence can be maximal.     

Predation promotes survival of beetles with lower resting metabolic rates

The energetic definition of fitness predicts that natural selection will maximize the residual energy available for growth and reproduction suggesting that energy metabolism might be a target of selection. In this experimental study, we investigated whether female and male yellow mealworm beetles, Tenebrio molitor L. (Coleoptera: Tenebrionidae), differ in their hiding behaviour, individual response latency time, and duration of immobility to treatments mimicking an approaching predation threat. We experimentally tested whether consistently repeatable anti‐predatory responses and resting metabolic rates (RMR) correlated with survival rates of individuals exposed to a nocturnal predator, the brown rat, Rattus norvegicus (Berkenhout) (Rodentia: Muridae). Resting metabolic rate was part of a syndrome involving anti‐predator behaviour. Individuals with lower RMR concealed themselves against predators in substrate more successfully than individuals with higher RMR, and hiding was associated with longer periods of immobility. Ultimately, mortality was higher in the high‐RMR beetles compared to the low‐RMR beetles. Our results provide direct evidence of natural selection against mobility, i.e., for reduced RMR in T. molitor beetles.     

Growth history and intrinsic factors influence risk assessment at a critical life transition for a fish

Making the appropriate decision in the face of predation risk dictates the fate of prey, and predation risk is highest at life history boundaries such as settlement. At the end of the larval phase, most coral reef fishes enter patches of reef containing novel predators. Since vision is often obscured in the complex surroundings, chemical information released from damaged conspecific is used to forewarn prey of an active predator. However, larvae enter the reef environment with their own feeding and growth histories, which will influence their motivation to feed and take risks. The present study explored the link between recent growth, feeding history, current performance and behavioural risk taking in newly settling stages of a coral reef damselfish (Pomacentrus amboinensis). Older and larger juveniles in good body condition had a stronger response to chemical alarm cues of injured conspecifics; these fish spent a longer time in shelter and displayed a more dramatic decrease in foraging behaviour than fish in lower body condition. Feeding experiments supported these findings and emphasized the importance of body condition in affecting risk assessment. Evidently, larval growth history and body condition influences the likelihood of taking risks under the threat of predation immediately after settlement, thereby affecting the probability of survival in P. amboinensis.     

锦鲫幼鱼标准代谢率与生长性能的关联

为考察温水性鲤科鱼类在不同食物资源条件下标准代谢率(Standard metabolic rate, SMR)与其生长性能的关系, 研究在(25.00.5)℃条件下测定体重相近的30尾锦鲫(Carassius auratus)幼鱼的SMR, 将所有实验鱼置于多单元格水槽进行实验处理(摄食与饥饿)。实验时间为4周(0、7d、14d、21d和28d), 包括两周的生长实验(014d)和两周的饥饿实验(1528d)。摄食期间, 于每天上午9: 00和下午21: 00用商业饲料对每尾鱼饱足投喂并记录投喂量。单尾鱼的体重和体长每隔1周测定一次, 而SMR仅在第0、第14和第28天测定。结果显示:(1)锦鲫幼鱼在摄食期间的体重、体长以及SMR均明显上升, 饥饿期间体重和SMR显著降低(P0.05), 但体长变化不明显; SMR不论摄食期间还是饥饿期间呈现较好的稳定性(二者P0.05)。(2)锦鲫幼鱼在摄食期间开始时的SMR (测定Ⅰ)与摄食率(FR)、摄食转化率(FE)以及特定体重生长率(SGRBM)均无相关性, 摄食期间结束时的SMR (测定Ⅱ)仅与FR呈正相关(P0.05), 但与FE以及SGRBM不相关。(3)摄食期间(饥饿期间)实验鱼的第一周日均体重增(减)量与第二周日均体重增(减)量呈正相关, 其体重的增率与减率无显著差异(P0.05)。(4)实验鱼在摄食期间的特定体重增长率(SGRBM)与饥饿期间的不相关(P0.05), 但摄食期间的特定SMR增长率(SGRSMR)与饥饿期间的呈负相关(P0.05)。研究表明在实验室环境中锦鲫幼鱼的SMR具有稳定性, 该种鱼的日均体重变化量保持相近, 此形态变化特征与SMR不相关, 并且该种鱼在实验起始时的SMR的个体差异未能预测其在实验室食物资源变动下生长性能的适应特征。     

Smell, learn and live: The role of chemical alarm cues in predator learning during early life history in a marine fish

The speed with which individuals can learn to identify and react appropriately to predation threats when transitioning to new life history stages and habitats will influence their survival. This study investigated the role of chemical alarm cues in both anti-predator responses and predator identification during a transitional period in a newly settled coral reef damselfish, Pomacentrus amboinensis. Individuals were tested for changes in seven behavioural traits in response to conspecific and heterospecific skin extracts. Additionally, we tested whether fish could learn to associate a previously novel chemical cue (i.e. simulated predator scent) with danger, after previously being exposed to a paired cue combining the conspecific skin extract with the novel scent. Fish exposed to conspecific skin extracts were found to significantly decreased their feeding rate whilst those exposed to heterospecific and control cues showed no change. Individuals were also able to associate a previously novel scent with danger after only a single previous exposure to the paired conspecific skin extract/novel scent cue. Our results indicate that chemical alarm cues play a large role in both threat detection and learned predator recognition during the early post-settlement period in coral reef fishes.     

Non‐pathogenic aquatic bacteria activate the immune system and increase predation risk in damselfly larvae

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Adaptive trade-offs in juvenile salmonid metabolism associated with habitat partitioning between coho salmon and steelhead trout in coastal streams

1. Adaptive trade-offs are fundamental to the evolution of diversity and the coexistence of similar taxa and occur when complimentary combinations of traits maximize efficiency of resource exploitation or survival at different points on environmental gradients. 2. Standard metabolic rate (SMR) is a key physiological trait that reflects adaptations to baseline metabolic performance, whereas active metabolism reflects adaptations to variable metabolic output associated with performance related to foraging, predator avoidance, aggressive interactions or migratory movements. Benefits of high SMR and active metabolism may change along a resource (productivity) gradient, indicating that a trade-off exists among active metabolism, resting metabolism and energy intake. 3. We measured and compared SMR, maximal metabolic rate (MMR), aerobic scope (AS), swim performance (UCrit) and growth of juvenile hatchery and wild steelhead and coho salmon held on high- and low-food rations in order to better understand the potential significance of variation in SMR to growth, differentiation between species, and patterns of habitat use along a productivity gradient. 4. We found that differences in SMR, MMR, AS, swim performance and growth rate between steelhead trout and coho salmon were reduced in hatchery-reared fish compared with wild fish. Wild steelhead had a higher MMR, AS, swim performance and growth rate than wild coho, but adaptations between species do not appear to involve differences in SMR or to trade-off increased growth rate against lower swim performance, as commonly observed for high-growth strains. Instead, we hypothesize that wild steelhead may be trading off higher growth rate for lower food consumption efficiency, similar to strategies adopted by anadromous vs. resident brook trout and Atlantic salmon vs. brook trout. This highlights potential differences in food consumption and digestion strategies as cryptic adaptations ecologically differentiating salmonid species. 5. We hypothesize that divergent digestive strategies, which are common and well documented among terrestrial vertebrates, may be an important but overlooked aspect of adaptive strategies of juvenile salmonids, and fish in general.     

Non-lethal effects of predation in birds

Predators can affect individual fitness and population and community processes through lethal effects (direct consumption or ‘density’ effects), where prey is consumed, or through non‐lethal effects (trait‐mediated effects or interactions), where behavioural compensation to predation risk occurs, such as animals avoiding areas of high predation risk. Studies of invertebrates, fish and amphibians have shown that non‐lethal effects may be larger than lethal effects in determining the behaviour, condition, density and distribution of animals over a range of trophic levels. Although non‐lethal effects have been well described in the behavioural ecology of birds (and also mammals) within the context of anti‐predation behaviour, their role relative to lethal effects is probably underestimated. Birds show many behavioural and physiological changes to reduce direct mortality from predation and these are likely to have negative effects on other aspects of their fitness and population dynamics, as well as affecting the ecology of their own prey and their predators. As a consequence, the effects of predation in birds are best measured by trade‐offs between maximizing instantaneous survival in the presence of predators and acquiring or maintaining resources for long‐term survival or reproduction. Because avoiding predation imposes foraging costs, and foraging behaviour is relatively easy to measure in birds, the foraging–predation risk trade‐off is probably an effective framework for understanding the importance of non‐lethal effects, and so the population and community effects of predation risk in birds and other animals. Using a trade‐off approach allows us to predict better how changes in predator density will impact on population and community dynamics, and how animals perceive and respond to predation risk, when non‐lethal effects decouple the relationship between predator density and direct mortality rate. The trade‐off approach also allows us to identify where predation risk is structuring communities because of avoidance of predators, even when this results in no observable direct mortality rate.     

Trade-Offs between Predation Risk and Growth Benefits in the Copepod Eurytemora affinis with Contrasting Pigmentation

Intraspecific variation in body pigmentation is an ecologically and evolutionary important trait; however, the pigmentation related trade-offs in marine zooplankton are poorly understood. We tested the effects of intrapopulation phenotypic variation in the pigmentation of the copepod Eurytemora affinis on predation risk, foraging, growth, metabolic activity and antioxidant capacity. Using pigmented and unpigmented specimens, we compared (1) predation and selectivity by the invertebrate predator Cercopagis pengoi, (2) feeding activity of the copepods measured as grazing rate in experiments and gut fluorescence in situ, (3) metabolic activity assayed as RNA:DNA ratio in both experimental and field-collected copepods, (4) reproductive output estimated as egg ratio in the population, and (5) total antioxidant capacity. Moreover, mitochondrial DNA (mtDNA) COI gene variation was analysed. The pigmented individuals were at higher predation risk as evidenced by significantly higher predation rate by C. pengoi on pigmented individuals and positive selection by the predator fed pigmented and unpigmented copepods in a mixture. However, the antioxidant capacity, RNA:DNA and egg ratio values were significantly higher in the pigmented copepods, whereas neither feeding rate nor gut fluorescence differed between the pigmented and unpigmented copepods. The phenotypic variation in pigmentation was not associated with any specific mtDNA genotype. Together, these results support the metabolic stimulation hypothesis to explain variation in E. affinis pigmentation, which translates into beneficial increase in growth via enhanced metabolism and antioxidant protective capacity, together with disadvantageous increase in predation risk. We also suggest an alternative mechanism for the metabolic stimulation via elevated antioxidant levels as a primary means of increasing metabolism without the increase in heat absorbance. The observed trade-offs are relevant to evolutionary mechanisms underlying plasticity and adaptation and have the capacity to modify strength of complex trophic interactions.     

宽鳍鱲个性与维持代谢、食物占有能力和日常能量消耗的关联

为了探讨集群性鱼类宽鳍鱲(Zacco platypus)的个性在群体中的生态学关联, 研究首先测定了宽鳍鱲的个性特征(包括活跃性、探索性、勇敢性和社会性)和标准代谢率。随后在群体(10个群体, 每群体6尾鱼)条件下测定了每尾鱼的运动轨迹、摄食状况, 并计算出每尾鱼的日总耗能。研究发现: 宽鳍鱲的个性特征与标准代谢率、摄食率和日总耗能均没有关联。个性与摄食率的关联丧失可能因为其野外生境食物丰度较高, 个性的差异未体现在对食物的竞争能力上; 而个性与能量代谢无关联可能与实验室单尾鱼测定条件下宽鳍鱲的应激反应有关。日总耗能与集群运动参数中的运动时间比、平均游泳速度和摄食率均呈正相关, 可能因为运动能耗和消化能耗在日总耗能中占比较高。另外, 日总耗能也与距质心距离正相关, 可能个性较积极的个体在群体中活跃程度较高, 集群时较多位于群体的边缘, 且日常能量消耗较高。研究结果表明, 集群时活跃的成员往往处于鱼群的外周, 食物占有能力更高, 但日常能量消耗也越多。研究还提示针对集群性野生鱼类, 可能在群体条件下考察其个性、能量代谢特征及其生态收益和代价更具生态学意义。     

Investment in defense and cost of predator-induced defense along a resource gradient

An organism’s investment in different traits to reduce predation is determined by the fitness benefit of the defense relative to the fitness costs associated with the allocation of time and resources to the defense. Inherent tradeoffs in time and resource allocation should result in differential investment in defense along a resource gradient, but competing models predict different patterns of investment. There are currently insufficient empirical data on changes in investment in defensive traits or their costs along resource gradients to differentiate between the competing allocation models. In this study, I exposed tadpoles to caged predators along a resource gradient in order to estimate investment in defense and costs of defense by assessing predator-induced plasticity. Induced defenses included increased tail depth, reduced feeding, and reduced swimming activity; costs associated with these defenses were reduced developmental rate, reduced growth, and reduced survival. At low resource availability, these costs predominately resulted in reduced survival, while at high resource availability the costs yielded a reduced developmental rate. Defensive traits responded strongly to predation risk, but did not respond to resource availability (with the exception of feeding activity), whereas traits construed as costs of defenses showed the opposite pattern. Therefore, defensive traits were highly sensitive to predation risk, while traits construed as costs of defense were highly sensitive to resource allocation tradeoffs. This difference in sensitivity between the two groups of traits may explain why the correlation between the expression of defensive traits and the expression of the associated defense costs was weak. Furthermore, my results indicate that genetic linkages and mechanistic integration of multiple defensive traits and their associated costs may constrain time and resource allocation in ways that are not addressed in existing models. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Compensation for predator-induced reduction in nestling feeding rate in the Great Spotted Woodpecker

Parent birds make efforts to prevent the immediate costs of predation through plastic behavioural responses to the actual predation risk, but this may incur future costs for offspring due to reduced parental care. However, the temporary nature of predator encounters suggests that nestling feeding reduced during the risky periods may be later compensated for by an increased feeding effort (the predation risk allocation hypothesis). We tested this prediction in the Great Spotted Woodpecker (Dendrocopos major) confronted with its major nest predator/competitor, the European Starling (Sturnus vulgaris). A brief encounter with a live starling was followed by a reduction in the nestling feeding rate, but the lost feedings were subsequently compensated for by an increased feeding rate. This compensatory effect was higher in older nestlings that are highly demanding in terms of energy requirements and fitness value from the parents’ perspective. Thus, birds are potentially able to respond not only to the immediate risk of nest predation by nest defence but also to compensate for the potential costs of nest defence in terms of unfulfilled nestling demands. However, data on the amount of delivered food are necessary to distinguish whether parents truly compensate for lost feeding or whether the increased feeding frequency represents a nest guarding strategy.     

Rainbow trout in seasonal environments: phenotypic trade‐offs across a gradient in winter duration

Survival through periods of resource scarcity depends on the balance between metabolic demands and energy storage. The opposing effects of predation and starvation mortality are predicted to result in trade‐offs between traits that optimize fitness during periods of resource plenty (e.g., during the growing season) and those that optimize fitness during periods of resource scarcity (e.g., during the winter). We conducted a common environment experiment with two genetically distinct strains of rainbow trout to investigate trade‐offs due to (1) the balance of growth and predation risk related to foraging rate during the growing season and (2) the allocation of energy to body size prior to the winter. Fry (age 0) from both strains were stocked into replicate natural lakes at low and high elevation that differed in winter duration (i.e., ice cover) by 59 days. Overwinter survival was lowest in the high‐elevation lakes for both strains. Activity rate and growth rate were highest at high elevation, but growing season survival did not differ between strains or between environments. Hence, we did not observe a trade‐off between growth and predation risk related to foraging rate. Growth rate also differed significantly between the strains across both environments, which suggests that growth rate is involved in local adaptation. There was not, however, a difference between strains or between environments in energy storage. Hence, we did not observe a trade‐off between growth and storage. Our findings suggest that intrinsic metabolic rate, which affects a trade‐off between growth rate and overwinter survival, may influence local adaptation in organisms that experience particularly harsh winter conditions (e.g., extended periods trapped beneath the ice in high‐elevation lakes) in some parts of their range.