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Trade-offs are central to life history theory and play a role in driving life history diversity. They arise from a finite amount of resources that need to be allocated among different functions by an organism. Yet covariation of demographic rates among individuals frequently do not reflect allocation trade-offs because of variation in resource acquisition. The covariation of traits among individuals can thus vary with the environment and often increases in benign environments. Surprisingly, little is known about how such context-dependent expression of trade-offs among individuals affect population dynamics across species with different life histories. To study their influence on population stability, we develop an individual-based simulation where covariation in demographic rates varies with the environment. We use it to simulate population dynamics for various life histories across the slow-fast pace-of-life continuum. We found that the population dynamics of slower life histories are relatively more sensitive to changes in covariation, regardless of the trade-off considered. Additionally, we found that the impact on population stability depends on which trade-off is considered, with opposite effects of intraindividual and intergenerational trade-offs. Last, the expression of different trade-offs can feed back to influence generation time through selection acting on individual heterogeneity within cohorts, ultimately affecting population dynamics. [ABSTRACT FROM AUTHOR]

An understanding of biological fitness is central to theory and practice in ecology and evolution, yet fitness remains an elusive concept to define and is challenging to measure accurately. Fitness reflects an individual's ability to pass its alleles on to subsequent generations. Researchers often quantify proxies for fitness, such as survival, growth, or reproductive success. However, it can be difficult to determine lifetime fitness, especially for species with long life spans. The abiotic and biotic environment strongly affects the expression of fitness, which means that fitness components can vary through both space and time. This spatial and temporal heterogeneity results in the impressive range of adaptations that we see in nature. Here, we review definitions of fitness and approaches to measuring fitness at the level of genes, individuals, genotypes, and populations and highlight that fitness is a key concept linking ecological and evolutionary thought. [ABSTRACT FROM AUTHOR]

How traits at multiple levels of biological organization evolve in a correlated fashion in response to directional selection is poorly understood, but two popular models are the very general "behavior evolves first" (BEF) hypothesis and the more specific "morphology-performance-behavior-fitness" (MPBF) paradigm. Both acknowledge that selection often acts relatively directly on behavior and that when behavior evolves, other traits will as well but most with some lag. However, this proposition is exceedingly difficult to test in nature. Therefore, we studied correlated responses in the high-runner (HR) mouse selection experiment, in which four replicate lines have been bred for voluntary wheel-running behavior and compared with four nonselected control (C) lines. We analyzed a wide range of traits measured at generations 20–24 (with a focus on new data from generation 22), coinciding with the point at which all HR lines were reaching selection limits (plateaus). Significance levels (226 P values) were compared across trait types by ANOVA, and we used the positive false discovery rate to control for multiple comparisons. This meta-analysis showed that, surprisingly, the measures of performance (including maximal oxygen consumption during forced exercise) showed no evidence of having diverged between the HR and C lines, nor did any of the life history traits (e.g., litter size), whereas body mass had responded (decreased) at least as strongly as wheel running. Overall, results suggest that the HR lines of mice had evolved primarily by changes in motivation rather than performance ability at the time they were reaching selection limits. In addition, neither the BEF model nor the MPBF model of hierarchical evolution provides a particularly good fit to the HR mouse selection experiment. [ABSTRACT FROM AUTHOR]

Trade-offs between life history traits are context dependent; they vary depending on environment and life stage. Negative associations between development and growth often characterize larval life stages. Both growth and development consume large parts of the energy budget of young animals. The metabolic rate of animals should reflect differences in growth and developmental rates. Growth and development can also have negative associations with immune function because of their costs. We investigated how intraspecific variation in growth and development affected the metabolism of larval amphibians and whether intraspecific variation in growth, development, and metabolic rate could predict mortality and viral load in larvae infected with ranavirus. We also compared the relationship between growth and development before and after infection with ranavirus. We hypothesized that growth and development would affect metabolism and predicted that each would have a positive correlation with metabolic rate. We further hypothesized that allocation toward growth and development would increase ranavirus susceptibility and therefore predicted that larvae with faster growth, faster development, and higher metabolic rates would be more likely to die from ranavirus and have higher viral loads. Finally, we predicted that growth rate and developmental rate would have a negative association. Intraspecific variation in growth rate and developmental rate did not affect metabolism. Growth rate, developmental rate, and metabolism did not predict mortality from ranavirus or viral load. Larvae infected with ranavirus exhibited a trade-off between developmental rate and growth rate that was absent in uninfected larvae. Our results indicate a cost of ranavirus infection that is potentially due to both the infection-induced anorexia and the cost of infection altering priority rules for resource allocation. [ABSTRACT FROM AUTHOR]

Premise of research. Cosexual plants reproduce as females by maturing seeds from fertilized ovules and as males by fertilizing their own ovules or those of other plants. Condition-dependent sex allocation occurs when the environment elicits quantitative differences in the rate at which fitness increases as resources are invested in male versus female function. Here, we examine whether resource availability and the pollination environment generate plastic responses in floral sex allocation and correlated traits. Methodology. We exposed the cosexual mustard species Brassica rapa to variation in resource availability (altered pot size) and in the pollination environment (unpollinated or fully pollinated). We assessed investment in anther and ovary length, which are known to correlate with pollen and ovule number; in flower buds produced at the onset of reproduction; and in buds produced approximately 15 d later. We also measured plant size and flower size, which are often correlated with increased allocation to female and male function, respectively. Pivotal results. In contrast to our predictions, anther-to-ovary length ratios at the onset of reproduction did not vary across plants of different sizes and were unaffected by the pot size treatment, despite an increase in anther lengths in the small pot treatment. Similarly, neither the pot size nor the pollination treatments elicited plasticity in anther-to-ovary length ratios over the course of reproduction. However, independent of plant size and as predicted, plants produced larger flowers over the course of reproduction in the pollination-absent treatment. Conclusions. We discuss the contexts in which resource availability and the pollination environment can influence sex allocation and provide cautionary advice on effective methods for experimentally generating and measuring sex allocation plasticity. [ABSTRACT FROM AUTHOR]

Premise of research. Inbreeding depression is well known to contribute to the maintenance of outcrossing in flowering plants because of its direct negative impacts on germination, survival, and fertility. Yet inbreeding depression could also impact fitness indirectly by affecting traits that govern biotic interactions. Relatively little is known about inbreeding depression on traits related to pollination success. Although often associated with pollinator attraction or reward, pollination success is also determined by floral longevity. Inbreeding depression in floral longevity has not been studied but has the potential to compound pollination disadvantages confronting inbred plants. Methodology. We investigated inbreeding depression for traits related to pollination success in the mixed-mating biennial Sabatia angularis L. (Pursh) (Gentianaceae). Under controlled, pollinator-free conditions, we performed crosses to create families of selfed and outcross seed and raised these plants to flower. We compared selfed and outcrossed individuals for the following traits: flower number, flower size, corolla longevity, stigma longevity, and plant height. We also considered whether inbreeding depression was influenced by flower number–longevity trade-offs or plant size. Pivotal results. Selfed plants produced significantly smaller flowers that lived for a significantly shorter amount of time than flowers of outcrossed plants, based on corolla and stigma longevity. Consistent with prior work, we found trade-offs between floral longevity and flower number and positive relationships between longevity and plant height. However, we did not detect inbreeding depression for flower number or plant size. Conclusions. We find novel evidence that inbreeding reduces both floral longevity and flower size. Although moderate, inbreeding depression for these traits could leave selfed plants at a disadvantage in pollen-limited environments, especially where rates of pollen export outpace deposition such that plants are forced to rely on pollinators for seed production. Our work contributes to our understanding of how inbreeding depression can alter plant-pollinator interaction outcomes and raises questions about its influence on reproductive allocation. [ABSTRACT FROM AUTHOR]

Prominent differences in aging among and within species present an evolutionary puzzle. The theories proposed to explain evolutionary differences in aging are based on the axiom that selection maximizes fitness, not necessarily life span. This implies trade-offs between investment in self-maintenance and investment in reproduction, where high investments in growth and current reproduction are associated with short life spans. Fast growth and large adult size are related to shorter life spans in the domestic dog, a bourgeoning model in aging research; however, whether reproduction influences life span in this system remains unknown. Here we test the relationship between reproduction and differences in life span among dog breeds, simultaneously controlling for shared ancestry and recent gene flow. We found that shared ancestry explains a higher proportion of the among-breed variation in life history traits, in comparison with recent gene flow. Our results also show that reproductive investment negatively impacts life span, and more strongly so in large breeds, an effect that is not merely a correlated response of adult size. These results suggest that basic life history trade-offs are apparent in a domestic animal whose diversity is the result of artificial selection and that among-breed differences in life span are due to a combination of size and reproduction. [ABSTRACT FROM AUTHOR]

Competition drives evolutionary change across taxa, but our understanding of how competitive differences among species directs the evolution of interspecific interactions remains incomplete. Verbal models assume that interspecific competition will select for reducing a species' sensitivity to competition with their opponent; however, they do not consider the potential for other demographic components of competitive ability to evolve, specifically, interspecific effects, intraspecific interactions, and intrinsic growth rates. To better understand how competitive ability evolves, we set out to explore how each component has evolved and whether their evolution has been constrained by trade-offs. By setting sympatric and allopatric populations of an annual grass in competition with a dominant invader, we demonstrate (1) that in response to interspecific competition, populations can evolve increased competitive ability through either reduced interspecific or, surprisingly, reduced intraspecific competition; (2) that trade-offs do not always constrain the evolution of competitive ability but rather that parameters may correlate in ways that mutually beget higher competitive ability; and (3) that the evolution of one species can influence the competitive ability of its opponent, a consequence of how competitive ability is defined ecologically. Overall, our results reveal the complexity with which demographic components evolve in response to interspecific competition and the impact past evolution can have on present-day interactions. [ABSTRACT FROM AUTHOR]

Traits often contribute to multiple functions, complicating our understanding of the selective pressures that influence trait evolution. In the Chihuahuan Desert, predation is thought to be the primary driver of cryptic light coloration in three White Sands lizard species relative to the darker coloration of populations on adjacent dark soils. However, coloration also influences radiation absorption and thus animal body temperatures. We combined comparative physiological experiments and biophysical models to test for thermal consequences of evolving different color morphs in White Sands across the three species. While light and dark morphs have not evolved different physiological heat limits within species, differences in radiation absorption between morphs lead to body temperature differences that impact relative overheating risk and activity patterns. Moreover, for all three species, an idealized morph that matches the White Sands substrate would have considerably less activity time, by approximately 1 month, than existing light morphs. Overall, there are both benefits and costs to greater substrate matching, the balance of which may prevent the evolution of optimal crypsis. Our work highlights the importance of color in dictating thermal performance and the complexity inherent in understanding the evolution of coloration. [ABSTRACT FROM AUTHOR]

Resource-intensive traits, such as dispersal and reproduction, can be difficult to express simultaneously because of resource limitations. One solution is to switch between resource-intensive behaviors. Such phenotypic plasticity is one strategy that organisms use to funnel resources from one expensive trait to another. In crickets (Gryllus texensis), the development and maintenance of flight muscles reduce resource availability for reproduction, leading to physiological trade-offs between the two traits. Long-winged female G. texensis can histolyze their wing muscles, resulting in increased egg production, but they can then no longer fly. Using a diet that mimics food availability in the field, we found that long-winged females adopted one of the three following strategies: early reproduction, intermediate reproduction, and late reproduction. Some late reproducers maintained their flight capability until the end of their natural life span and laid few eggs. If females lost the ability to fly (i.e., their hind wings are removed), they laid eggs earlier, leading to increased reproductive output. However, other environmental cues (e.g., an increased number of mates, increased oviposition substrate quality, or a bout of dispersal flight) had no effect. Late-reproducing females laid 96% fewer eggs than early reproducers, suggesting that late reproduction exacts a huge fitness cost. Nevertheless, some females maintain their flight muscles to the end of their natural life span in both the lab and the field. We suggest that the ability to fly allows for bet hedging against an environmental catastrophe (e.g., drought or flood). This benefit may help explain the persistence of late reproducing long-winged females, despite the cost of this choice. As climate change increases drought and flood in Texas, late dispersal may be one factor that helps this species survive in the future. An increased understanding of factors that maintain seemingly low fitness strategies can help us predict the resilience of species under climate change. [ABSTRACT FROM AUTHOR]

While species distribution models (SDMs) are widely used to predict the vulnerability of species to climate change, they do not explicitly indicate the extent to which plastic responses ameliorate climate change impacts. Here we use data on plastic responses of 32 species of Drosophila to desiccation stress to suggest that basal resistance, rather than adult hardening, is relatively more important in determining species differences in desiccation resistance and sensitivity to climate change. We go on to show, using the semimechanistic SDM CLIMEX, that the inclusion of plasticity has some impact on current species distributions and future vulnerability for widespread species but has little impact on the distribution of arguably more vulnerable tropically restricted species. [ABSTRACT FROM AUTHOR]

Consumers regularly experience trade-offs in their ability to find, handle, and digest different resources. Evolutionary ecologists recognized the significance of this observation for the evolution and maintenance of biological diversity long ago and continue to elaborate on the conditions under which to expect one or several specialists, generalists, or combinations thereof. Existing theory based on a single evolving trait predicts that specialization requires strong trade-offs such that generalists perform relatively poorly, while weak trade-offs favor a single generalist. Here, we show that this simple dichotomy does not hold true under joint evolution of two or more foraging traits. In this case, the boundary between trade-offs resulting in resource specialists and resource generalists is shifted toward weaker trade-off curvatures. In particular, weak trade-offs can result in evolutionary branching, leading to the evolution of two coexisting resource specialists, while the evolution of a single resource generalist requires particularly weak trade-offs. These findings are explained by performance benefits due to epistatic trait interactions enjoyed by phenotypes that are specialized in more than one trait for the same resource. [ABSTRACT FROM AUTHOR]

Many species facing climate change have complex life cycles, with individuals in different stages differing in their sensitivity to a changing climate and their contribution to population growth. We use a quantitative genetics model to predict the dynamics of adaptation in a stage-structured population confronted with a steadily changing environment. Our model assumes that different optimal phenotypic values maximize different fitness components, consistent with many empirical observations. In a constant environment, the population evolves toward an equilibrium phenotype, which represents the best compromise given the trade-off between vital rates. In a changing environment, however, the mean phenotype in the population will lag behind this optimal compromise. We show that this lag may result in a shift along the trade-off between vital rates, with negative consequences for some fitness components but, less intuitively, improvements in some others. Complex eco-evolutionary dynamics can emerge in our model due to feedbacks between population demography and adaptation. Because of such feedback loops, selection may favor further shifts in life history in the same direction as those caused by maladaptive lags. These shifts in life history could be wrongly interpreted as adaptations to the new environment, while in reality they only reflect the inability of the population to adapt fast enough. [ABSTRACT FROM AUTHOR]

Predicting the evolution of expanding populations is critical to controlling biological threats such as invasive species and cancer metastasis. Expansion is primarily driven by reproduction and dispersal, but nature abounds with examples of evolution where organisms pay a reproductive cost to disperse faster. When does selection favor this "survival of the fastest"? We searched for a simple rule, motivated by evolution experiments where swarming bacteria evolved into a hyperswarmer mutant that disperses ∼100% faster but pays a growth cost of ∼10% to make many copies of its flagellum. We analyzed a two-species model based on the Fisher equation to explain this observation: the population expansion rate (v) results from an interplay of growth (r) and dispersal (D) and is independent of the carrying capacity: v = 2 (rD) 1 / 2 . A mutant can take over the edge only if its expansion rate (v 2) exceeds the expansion rate of the established species (v 1); this simple condition ( v 2 > v 1 ) determines the maximum cost in slower growth that a faster mutant can pay and still be able to take over. Numerical simulations and time-course experiments where we tracked evolution by imaging bacteria suggest that our findings are general: less favorable conditions delay but do not entirely prevent the success of the fastest. Thus, the expansion rate defines a traveling wave fitness, which could be combined with trade-offs to predict evolution of expanding populations. [ABSTRACT FROM AUTHOR]

High predation risk can favor rapid offspring development at the expense of offspring quality. Impacts of rapid development on phenotypic quality should be most readily expressed in traits that minimize fitness costs. We hypothesize that ephemeral traits that are replaced or repaired after a short period of life might express trade-offs in quality as a result of rapid development more strongly than traits used throughout life. We explored this idea for plumage quality in nestling body feathers, an ephemeral trait. We found a strong trade-off whereby nestlings that spend less time in the nest produced lower-quality plumage with less dense barbs relative to adults across 123 temperate and tropical species. For a subset of these species (n = 67), we found that variation in the risk of nest predation explained additional variation in plumage quality beyond development time. Ultimately, the fitness costs of a poor-quality ephemeral trait, such as nestling body feathers, may be outweighed by the fitness benefits of shorter development times that reduce predation risk. At the same time, reduced resource allocation to traits with small fitness costs, such as ephemeral traits, may ameliorate resource constraints from rapid development on traits with larger fitness impacts. [ABSTRACT FROM AUTHOR]

Premise of research. Low-dose ionizing radiation (LDIR) can generate deleterious mutations that may have significant negative impact on life-history traits. Methodology. Here we quantified the germination rate of seeds of 25 species of plants from multiple sampling locations at Chernobyl when grown under similar (controlled, nonirradiated, common garden) greenhouse conditions. Pivotal results. We found a significant negative relationship between the germination rate of seeds in an uncontaminated common garden and ambient radiation in the field from which seeds were collected with an overall intermediate effect size. Although there was significant variation in the effect of radiation on germination rate among taxa, we found no significant effect of radiation due to genome size, number of chromosomes, level of ploidy, or seeds that originated from pollination by bi- or trinucleate pollen grains. Seed weight was significantly reduced in sites with higher radiation (0.03-129.1 µSv/h). There was a significant effect of radiation by seed weight interaction on germination rate. Conclusions. The effect of LDIR on pollen viability was positively correlated with the effect of radiation on germination rate, suggesting that a common underlying factor, most probably deleterious mutations, accounted for this common negative impact of radiation on both pollen viability and germination rate. [ABSTRACT FROM AUTHOR]

The environment can affect individual performance directly via resource availability or indirectly through resource allocation among competing fitness components, such as body growth and maintenance activities related to short-term survival. Corticosterone (CORT), the main glucocorticoid in birds, may be an important mediator of energy allocation to different organismal functions, but its effect on the plasticity of fitness-related traits has rarely been investigated at different ontogenetic stages. Here, we evaluated the role of baseline and stress-induced CORT on nestling development of wild great tits (Parus major) under different growth conditions and at different developmental stages. We found that nestling mass and condition were positively related to stress-induced levels of total CORT and free CORT in the middle of nestling stage but negatively related at the prefledging stage. Our results also showed that stress-induced levels of total CORT can be used as a proxy for bioavailable free CORT. No significant relationship of body mass or condition in either stage was shown with levels of baseline CORT or stress-induced CORT-binding globulin capacity. None of these age-specific relationships depend on brood size manipulation. We conclude that the effects of glucocorticoids on nestling performance are stage specific. The ability to secrete CORT may have beneficial effects on nestling development in the middle of the nestling stage, while it can be adaptive to keep CORT levels at optimal ranges before fledging to give priority to growth-related processes that are important for long-term fitness. [ABSTRACT FROM AUTHOR]

One of the central tenets of life-history theory is that organisms cannot simultaneously maximize all fitness components. This results in the fundamental trade-off between reproduction and life span known from numerous animals, including humans. Social insects are a well-known exception to this rule: reproductive queens outlive nonreproductive workers. Here, we take a step forward and show that under identical social and environmental conditions the fecundity-longevity trade-off is absent also within the queen caste. A change in reproduction did not alter life expectancy, and even a strong enforced increase in reproductive efforts did not reduce residual life span. Generally, egg-laying rate and life span were positively correlated. Queens of perennial social insects thus seem to maximize at the same time two fitness parameters that are normally negatively correlated. Even though they are not immortal, they best approach a hypothetical "Darwinian demon" in the animal kingdom. [ABSTRACT FROM AUTHOR]

The parchment-shelled eggs of squamate reptiles take up substantial water from the nest environment, enabling the conversion of yolk into neonatal tissue and buffering the embryo against the possibility of subsequent dry weather. During development, increasing amounts of water are stored in the embryonic sacs (i.e., membranes around the embryo: amnion, allantois, and chorion). The evolution of viviparity (prolonged uterine retention of developing embryos) means that embryonic-sac fluid storage now imposes a cost (increased maternal burdening), confers less benefit (because the mother buffers fetal water balance), and introduces a potential conflict among uterine siblings (for access to finite water supplies). Our data on nine species of squamate reptiles and published information on three species show that the embryonic-sac fluids comprise around 33% of neonatal mass in viviparous species versus 94% in full-term eggs of oviparous squamates. Data on parturition in 149 vipers (Vipera aspis, a viviparous species) show that larger offspring store more fluids in their fetal sacs and that an increase in litter size is associated with a decrease in fluid-sac mass per offspring. Overall, the evolutionary transition from oviparity to viviparity may have substantially altered selective forces on offspring packaging and created competition among offspring for access to water reserves during embryonic development. [ABSTRACT FROM AUTHOR]

Brain size differs substantially among species, and several hypotheses have been proposed to explain the evolution of brain size. Because the brain is among themost energetically expensive organs in the vertebrate body, trade-offs have been hypothesized to exert constraints on brain size evolution. Prominently, the expensive tissue hypothesis (ETH) proposes that reducing the size of another expensive organ, such as the gut, should compensate for the cost of a large brain. But energetic constraintsmay also drive covariation between the brain and other costly traits--such as bodymaintenance, locomotion, or reproduction-- as formulated in the energy trade-off hypothesis. To date, these hypotheses have mainly been tested in homeothermic animals and within the ectothermic animals, primarily in fishes. Here, we undertake a comparative test of the interplay between energetic limitations and brain size evolution within amphibians. After controlling for phylogenetic relationships and body size, we find a negative correlation between brain mass and the length of the digestive tract within 30 species of anurans. We further find that the evolution of large brain size is accompanied by an increase in female reproductive investment into egg size. Our results suggest that the evolution of brain size follows general patterns across vertebrate clades. [ABSTRACT FROM AUTHOR]

Yolk androgens of maternal origin are important mediators of prenatal maternal effects. Although in many species short-term benefits of exposure to high yolk androgen concentrations for the offspring have been observed, females differ substantially in the amount of androgens they transfer to their eggs. It suggests that costs for the offspring or the mother constrain the evolution of maternal hormone transfer. However, to date, the nature of these costs remains poorly understood. Unlike most previous work that focused on potential costs for the offspring, we here investigated whether high yolk testosterone transfer is associated with metabolic costs (i.e., a higher metabolic rate) for the mother. We show that Japanese quail (Coturnix japonica) females that deposit higher testosterone concentrations into their eggs have a higher resting metabolic rate. Because a higher metabolic rate is often associated with a shorter life span, this relationship may explain the negative association between yolk testosterone transfer and female longevity observed in the wild. Our results suggest that metabolic costs for themothercan balance the short-term benefits of yolk testosterone exposure for the offspring, thereby contributing to the maintenance of variation in maternal yolk hormone transfer in natural populations. [ABSTRACT FROM AUTHOR]

Life-history theory predicts that organisms will hedge current reproductive investment against potential costs in terms of survivorship and future fecundity. However, little is known regarding the endocrine mechanisms underlying bet-hedging strategies in free-ranging male vertebrates. We examined the relationships among individual energetic status, steroid hormones, mate search, and reproductive behavior in free-ranging male timber rattlesnakes. Snakes were monitored over four active seasons in order to test two hypotheses: (1) males adjust the amount of time and energy allocated toward reproduction according to the level of individual energy stores, and (2) observed condition-dependent reproductive allocation is associated with circulating concentrations of steroid hormones (testosterone and corticosterone) thought to regulate reproductive behaviors in vertebrates. A positive relationship between body condition and testosterone was observed in both the field and the laboratory. Male mate search effort was positively correlated with both body condition and testosterone. Body condition and testosterone concentrations were negatively related to time allocated toward foraging during the breeding season. A strong effect of year was observed in the analysis of testosterone and search effort, suggesting that multiple environmental factors impact hormone production and reproductive investment. Corticosterone was not related to any measured variable. Therefore, our results did not indicate a clear role of corticosterone in mediating observed relationships between energetic status and behavior. Observed relationships are consistent with the hypothesis that males allocate time and energy toward reproduction according to individual energetic status and that testosterone plays a role in mediating the trade-off between current reproductive investment and residual reproductive value. [ABSTRACT FROM AUTHOR]

Different mechanisms have been proposed to explain how vertical and horizontal heterogeneity in light conditions enhances tree species coexistence in forest ecosystems. The foliage partitioning theory proposes that differentiation in vertical foliage distribution, caused by an interspecific variation in mortality-to-growth ratio, promotes stable coexistence. In contrast, successional niche theory posits that horizontal light heterogeneity, caused by gap dynamics, enhances species coexistence through an interspecific trade-off between growth rate and survival. To distinguish between these theories of species coexistence, we analyzed tree inventory data for 370 species from the 50-ha plot in Pasoh Forest Reserve, Malaysia. We used community-wide Bayesian models to quantify size-dependent growth rate and mortality of every species. We compared the observed size distributions and the projected distributions from size-dependent demographic rates. We found that the observed size distributions were not simply correlated with the rate of population increase but were related to demographic properties such as size growth rate and mortality. Species with low relative abundance of juveniles in size distribution showed high growth rate and low mortality at small tree sizes and low per-capita recruitment rate. Overall, our findings were in accordance with those predicted by foliage partitioning theory. [ABSTRACT FROM AUTHOR]

We explore pathogen virulence evolution during the spatial expansion of an infectious disease epidemic in the presence of a novel host movement trade-off, using a simple, spatially explicit mathematical model. This work is motivated by empirical observations of the Mycoplasma gallisepticum invasion into North American house finch (Haemorhous mexicanus) populations; however, our results likely have important applications to other emerging infectious diseases in mobile hosts. We assume that infection reduces host movement and survival and that across pathogen strains the severity of these reductions increases with pathogen infectiousness. Assuming these trade-offs between pathogen virulence (host mortality), pathogen transmission, and host movement, we find that pathogen virulence levels near the epidemic front (that maximize wave speed) are lower than those that have a short-term growth rate advantage or that ultimately prevail (i.e., are evolutionarily stable) near the epicenter and where infection becomes endemic (i.e., that maximize the pathogen basic reproductive ratio). We predict that, under these trade-offs, less virulent pathogen strains will dominate the periphery of an epidemic and that more virulent strains will increase in frequency after invasion where disease is endemic. These results have important implications for observing and interpreting spatiotemporal epidemic data and may help explain transient virulence dynamics of emerging infectious diseases. [ABSTRACT FROM AUTHOR]

Although little studied in natural populations, the persistence of immuno-globulins may dramatically affect the dynamics of immunity and the ecology and evolution of host-pathogen interactions involving vertebrate hosts. By means of a multiple-year vaccination design against Newcastle disease virus, we experimentally addressed whether levels of specific antibodies can persist over several years in females of a long-lived procellariiform seabird-Cory's shearwater-and whether maternal antibodies against that antigen could persist over a long period in offspring several years after the mother was exposed. We found that a single vaccination led to high levels of antibodies for several years and that the females transmitted antibodies to their offspring that persisted for several weeks after hatching even 5 years after a single vaccination. The temporal persistence of maternally transferred antibodies in nestlings was highly dependent on the level at hatching. A second vaccination boosted efficiently the level of antibodies in females and thus their transfer to offspring. Overall, these results stress the need to consider the temporal dynamics of immune responses if we are to understand the evolutionary ecology of host-parasite interactions and trade-offs between immunity and other life-history characteristics, in particular in long-lived species. They also have strong implications for conservation when vaccination may be used in natural populations facing disease threats. [ABSTRACT FROM AUTHOR]

Hydroperiod strongly influences the breeding period and development time of many amphibians, and larvae of various species display developmental plasticity in response to habitat drying. Hydrologic alterations associated with anthropogenic activities potentially can influence host--parasite interactions in humans and wildlife, but few investigators have examined this possibility. Pathogens can delay amphibian development, so infections could constrain the ability of parasitized larvae to respond to a shortened hydroperiod under drying conditions. We examined the individual and joint effects of hydroperiod and infection with a pathogenic trematode parasite (Ribeiroia ondatrae) on larval development of Pacific chorus frogs (Pseudacris regilla) in mesocosms. Tadpoles subjected to accelerated drying were twice as likely to emerge early. However, this tendency was most pronounced among uninfected individuals (96% of early-emerging individuals) and may have arisen from differences in competitive ability between infected and uninfected conspecifics in mesocosms. Our findings indicate the importance of investigating hydrologic variables associated with environmental perturbations, such as climate change, particularly as they relate to the ability of organisms to respond to multiple stressors that include diseases. [ABSTRACT FROM AUTHOR]

Premise of research. Sex allocation theory considers that, given limited resources, a trade-off must exist between allocations to competing functions, such as accessory and primary sexual organs, or between allocations to primary male and female sexual organs. However, phenotypic and genotypic correlations between floral components have often been found to be positive. A view of resource allocation from a phenotypic-integration perspective could shed light on these apparently paradoxical results. Resource allocation, and not only floral metric traits, can also be signi?cantly integrated because of genetic or developmental constraints or functional contribution to offspring production. We describe the allocation of resources to inflorescence components and assess for the first time whether resources can also be phenotypically integrated in Tussilago farfara L. (Asteraceae), a species with inflorescences functioning as pollination units. Methodology. We studied the absolute and proportional allocation of resources to inflorescence components in terms of dry mass, N, and P as well as their nutrient concentration.We applied indices of phenotypic integration, previously limited to metric traits, to resource allocation at the inflorescence level. Pivotal results. Irrespective of currency and site differences, (1) scapes were always the inflorescence components receiving the highest allocation, whereas rays received the lowest allocation; and (2) allocation to inflorescence components was phenotypically integrated, ranging from 2.80% to 58.44% of the maximum possible integration. Conclusions. Integration in terms of resources was similar to or higher than that reported for floral metric traits in other species. The phenotypic integration of resources might result from the fact thatmost inflorescence components of T. farfara may contribute significantly to different functions, such as support, pollen donation, and seed production. The generality of the integration of resources allocated to sexual reproductive components awaits future studies but might help in understanding the usual lack of negative correlations between floral components expected to be linked by trade-offs. [ABSTRACT FROM AUTHOR]

One key trade-off underlying life-history evolution is the one between age and size at maturity, with earlier maturation leading to greater chances of juvenile survival at the cost of reduced fecundity as an adult. Here we model the impact of limited dispersal and kin competition on the stable resolution of this trade-off. We show that if mating is at least occasionally nonlocal, then limited dispersal favors juvenile survival over adult fecundity in females, promoting earlier female maturation at the population level; at the same time, it favors adult fecundity over juvenile survival in males, promoting later male maturation. Limited dispersal and local competition can thus drive the evolution of sexual dimorphism in the timing of maturation and consequent dimorphism in body size. At the individual level, if maturation can be flexibly adjusted in response to dispersal status, then both males and females who disperse as offspring should mature earlier than those who remain on their natal patch. [ABSTRACT FROM AUTHOR]

Studies of trilled vocalizations provide a premiere illustration of how performance constraints shape the evolution of mating displays. In trill production, vocal tract mechanics impose a tradeoff between syllable repetition rate and frequency bandwidth, with the trade-off most pronounced at higher values of both parameters. Available evidence suggests that trills that simultaneously maximize both traits are more threatening to males or more attractive to females, consistent with a history of sexual selection favoring highperformance trills. Here, we identify a sampling limitation that confounds the detection and description of performance trade-offs. We reassess 70 data sets (from 26 published studies) and show that sampling limitations afflict 63 of these to some degree. Traditional upper-bound regression, which does not control for sampling limitations, detects performance trade-offs in 33 data sets; yet when sampling limitations are controlled, performance trade-offs are detected in only 15. Sampling limitations therefore confound more than half of all performance trade-offs reported using the traditional method. An alternative method that circumvents this sampling limitation, which we explore here, is quantile regression. Our goal is not to question the presence of mechanical trade-offs on trill production but rather to reconsider how these trade-offs can be detected and characterized from acoustic data. [ABSTRACT FROM AUTHOR]

The use of species traits in basic and applied ecology is expanding rapidly because trait-based approaches hold the promise to increase our mechanistic understanding of biological responses. Such understanding could transform descriptive field studies in community ecology into predictive studies. Currently, however, trait-based approaches often fail to reflect species-environment relationships adequately. The difficulties have been perceived mainly as methodological, but we suggest that the problem is more profound and touches on the fundamentals of ecology and evolution. Selection pressures do not act independently on single traits, but rather, on species whose success in a particular environment is controlled by many interacting traits. Therefore, the adaptive value of a particular trait may differ across species, depending on the other traits possessed by the species and the constraints of its body plan. Because of this context-dependence, trait-based approaches should take into account the way combinations of traits interact and are constrained within a species. We present a new framework in which trade-offs and other interactions between biological traits are taken as a starting point from which to develop a better mechanistic understanding of species occurrences. The framework consists of 4 levels: traits, trait interactions, trait combinations, and life-history strategies, in a hierarchy in which each level provides the building blocks for the next. Researchers can contribute knowledge and insights at each level, and their contributions can be verified or falsified using logic, theory, and empirical data. Such an integrated and transparent framework can help fulfill the promise of traits to transform community ecology into a predictive science. [ABSTRACT FROM AUTHOR]

The metabolic theory of ecology (MTE) states that metabolic rate, ruled mainly by individual mass and temperature, determines many other biological rates. This view of ecology as ruled by the laws of physics and thermodynamics contrasts with life-history-optimization (LHO) theories, where traits are shaped by evolutionary processes. Integrating the MTE and LHO can lead, however, to a synthetic theory of ecology. In this work, we link the two theories to show that offspring development time is the result of both maternal investment in offspring and the metabolic constraints on offspring growth. We formulate a model that captures how offspring development time is the consequence of both offspring growth rate, determined by temperature and allometric scaling in accordance with the MTE, and the size reached by offspring at the end of the developmental period, determined mainly by LHO and reproductive strategies. We first extend the trade-off between offspring size and offspring number to ectotherms, showing that increased body temperatures result in increased resources available for reproduction. We then combine this trade-off with the general ontogenetic growth model to show that there is a trade-off between the number of offspring produced and offspring development time. The model predicts a shorter developmental time in organisms producing larger numbers of offspring. [ABSTRACT FROM AUTHOR]

The exact nature of the relationship among species range sizes, speciation, and extinction events is not well understood. The factors that promote larger ranges, such as broad niche widths and high dispersal abilities, could increase the likelihood of encountering new habitats but also prevent local adaptation due to high gene flow. Similarly, low dispersal abilities or narrower niche widths could cause populations to be isolated, but such populations may lack advantageous mutations due to low population sizes. Here we present a large-scale, spatially explicit, individual-based model addressing the relationships between species ranges, speciation, and extinction. We followed the evolutionary dynamics of hundreds of thousands of diploid individuals for 200,000 generations. Individuals adapted to multiple resources and formed ecological species in a multidimensional trait space. These species varied in niche widths, and we observed the coexistence of generalists and specialists on a few resources. Our model shows that species ranges correlate with dispersal abilities but do not change with the strength of fitness trade-offs; however, high dispersal abilities and low resource utilization costs, which favored broad niche widths, have a strong negative effect on speciation rates. An unexpected result of our model is the strong effect of underlying resource distributions on speciation: in highly fragmented landscapes, speciation rates are reduced. [ABSTRACT FROM AUTHOR]

All organisms must trade off resource allocation between different life processes that determine their survival and reproduction. Malaria parasites replicate asexually in the host but must produce sexual stages to transmit between hosts. Because different specialized stages are required for these functions, the division of resources between these life-history components is a key problem for natural selection to solve. Despite the medical and economic importance of these parasites, their reproductive strategies remain poorly understood and often seem counterintuitive. Here, we tested recent theory predicting that in-host competition shapes how parasites trade off investment in in-host replication relative to between- host transmission. We demonstrate, across several genotypes, that Plasmodium chabaudi parasites detect the presence of competing genotypes and facultatively respond by reducing their investment in sexual stages in the manner predicted to maximize their competitive ability. Furthermore, we show that genotypes adjust their allocation to sexual stages in line with the availability of exploitable red blood cell resources. Our findings are predicted by evolutionary theory developed to explain life-history trade-offs in more traditionally studied multicellular taxa and suggest that the answer to the long-standing question of why so few transmission stages are produced is that in most natural infections heavy investment in reproduction may compromise in-host survival. [ABSTRACT FROM AUTHOR]

"The jack-of-all-trades is a master of none" describes the widely held belief that engaging in many tasks comes at the cost of being unable to do those tasks well. However, empirical evidence for generalist fitness costs remains scarce. We used published data from a long-term field survey of aphid parasitoids to determine whether relative specialists are more abundant than generalists on their shared hosts, a pattern that would be expected if generalists suffer a trade-off between host-range breadth and host-use efficiency. Relative specialists were more abundant than generalists on their shared hosts, but only when we used a measure of specialization that accounts for the taxonomic differences among parasitoids' hosts. These results suggest that a generalist-specialist trade-off exists within this group of parasitoids and that the generalist fitness cost depends on the taxonomic breadth, rather than the number, of host species that are used. [ABSTRACT FROM AUTHOR]

Although temporal variability in the physical environment plays a major role in population fluctuations, little is known about how it drives the ecological and evolutionary dynamics of species interactions. We studied experimentally how extrinsic re- source pulses affect evolutionary and ecological dynamics between the prey bacterium Serratia marcescens and the predatory protozoan Tetrahymena thermophila. Predation increased the frequency of defensive, nonpigmented prey types, which bore competitive costs in terms of reduced maximum growth rate, most in a constant-resource environment. Furthermore, the predator densities of the pulsed- resource environment regularly fluctuated above and below the mean predator densities of the constant environment. These results suggest that selection favored fast-growing competitor prey types over defensive but slower-growing prey types more often in the pulsed-resource environment (abundance of resources and low predation risk). As a result, the selection for prey defense fluctuated more in the pulsed-resource environment, leading to a weaker mean response in prey defense. At the ecological level, the evolution of prey defense weakened the relative strength of top-down regulation on prey community. This was more evident in the constant-resource environment, whereas the slow emergence of defensive prey types gradually decreased the amplitude of predator peaks in the pulsed-resource environment. Our study suggests that rapid evolution plays a smaller role in the ecological dynamics of communities dominated by resource pulses. [ABSTRACT FROM AUTHOR]

The experience of environmental stress during development can substantially affect an organism's life history. These effects are often mainly negative, but a growing number of studies suggest that under certain environmental conditions early experience of such stress may yield individuals that are less sensitive to environmental stress later on in life. We used the butterfly Bicyclus anynana to study the effects of limited larval and adult food and forced flight on individual performance measured as reproduction and adult life span. Larvae exposed to food stress showed longer development and produced smaller adults. Thus, they were not able to fully compensate for the food deprivation during development. Females that experienced food stress during development did not increase tolerance for adult food limitation. However, females exposed to food stress during development coped better with forced flight compared with the control group. The apparent absence of costs of flight in poor-quality females may be a by-product of an altered body allocation, as females experiencing both food stress treatments had increased thorax ratios, compared with controls, and increased flight performances. The results reveal an important plasticity component to variation in flight performance and suggest that the cost of flight depends on an individual's internal condition. [ABSTRACT FROM AUTHOR]

Bet hedging is a means to increase fitness in environments that vary unpredictably in space and time. In such environments, models predict a trade-off between the bet-hedging strategies dispersal and dormancy, while the increasing importance of risk reduction with decreasing predictability should lead to an increase in dispersal and dormancy along gradients of environmental predictability. However, so far there has been no experimental study to test these predictions in the field. Here, we used a set of novel field experiments that enabled us to quantify and separate seedling recruitment from three sources: local reproduction, dormancy, and dispersal. The study included the entire plant community from five environments differing considerably in predictability. Evidence for both the existence of a trade-off between dispersal and dormancy within environments and their increased use in unpredictable environments was very weak. The importance of dispersal for population and community dynamics in our system was extremely low relative to dormancy and local reproduction. This indicates that the role of dispersal for buffering environmental variation may be negligible compared with other risk-reducing strategies. Our findings highlight the urgent need for multispecies and multisite experiments in empirical tests of theoretical predictions. [ABSTRACT FROM AUTHOR]

Insect herbivory is a major driving force of plant evolution. Phenotypic plasticity and developmental variation provide a means for plants to cope with variable herbivory. We characterized the genetics of developmental variation and phenotypic plasticity in trichome density, a putative defensive trait of Mimulus guttatus (yellow monkeyflower). Our results are evaluated in relation to the optimal defense theory, which provides testable predictions for plastic and developmental patterns in defense traits. We found that both developmental stage and simulated insect damage affected trichome production, but in different ways. Plants were more likely to produce at least some trichomes on later leaves than on earlier leaves, regardless of damage. Damage did not affect the average probability of producing trichomes, but it did increase the density of hairs on trichome-positive plants. We mapped trichome quantitative trait loci (QTL) by selectively genotyping a large panel of recombinant inbred lines derived from two highly divergent populations. Several highly pleiotropic QTL influenced multiple aspects of the trichome phenotype (constitutive, developmental, and/or plastic responses). Only one of the QTL influenced trichome induction following damage. In a result that is consistent with a central prediction of optimal defense theory, the high allele at this location was from the ancestral population with low constitutive trichome production. [ABSTRACT FROM AUTHOR]

We evaluate the cost-of-reproduction hypothesis in the burying beetle Nicrophorus orbicollis and examine how the importance of this trade-off changes as females age (i.e., the terminal-investment hypothesis). These beetles breed on small vertebrate carcasses, which serve as a food resource for them and their offspring. Consistent with the cost-of-reproduction hypothesis, females manipulated to overproduce offspring suffered a reduction in fecundity and life span when compared to controls, although all reproducing females had reduced life spans compared to nonbreeding females. Older females produced larger broods and allocated less of the carcass to their own body mass and a greater proportion to offspring than did younger females. Resource allocation to offspring increased with age. Females given larger carcasses invested more in current reproduction and less in future reproduction than did females given smaller carcasses. Our results provide unconfounded support for both the cost-of-reproduction hypothesis (i.e., current reproduction constrains future reproductive output) and the terminal-investment hypothesis (i.e., the importance of the trade-off between current and future reproduction declines with age such that allocation to current reproduction should increase as females age). [ABSTRACT FROM AUTHOR]

We derive a comprehensive overview of specialization evolution based on analytical results and numerical illustrations. We study the separate and joint evolution of two critical facets of specialization- local adaptation and habitat choice-under different life cycles, modes of density regulation, variance-covariance structures, and trade-off strengths. A particular feature of our analysis is the investigation of arbitrary trade-off functions. We find that local-adaptation evolution qualitatively changes the outcome of habitat-choice evolution under a wide range of conditions. In addition, habitat-choice evolution qualitatively and invariably changes the outcomes of local-adaptation evolution whenever trade-offs are weak. Even weak trade-offs, which favor generalists when habitat choice is fixed, select for specialists once local adaptation and habitat choice are both allowed to evolve. Unless trapped by maladaptive genetic constraints, joint evolution of local adaptation and habitat choice in the models analyzed here thus always leads to specialists, independent of life cycle, density regulation, and trade-off strength, thus raising the bar for evolutionarily sound explanations of generalism. Whether a single specialist or two specialists evolve depends on the life cycle and the mode of density regulation. Finally, we explain why the gradual evolutionary emergence of coexisting specialists requires more restrictive conditions than does their evolutionarily stable maintenance. [ABSTRACT FROM AUTHOR]

For ectotherms, growth and reproduction are possible only during a limited period of the year in seasonal environments. In insects, fitness is generally maximized by producing as many generations as possible within a season, and in many species, the number of generations produced (voltinism) increases with increasing season length. In this study, we analyzed variation in adult life histories in insects along a climatic gradient. The analyzed trait is reproductive effort (resource allocation to reproduction). We begin by formalizing the trade-off between current reproduction and subsequent survival generated by reproductive effort. It appeared that reproductive effort is correlated positively with early fecundity and negatively with lifetime fecundity and life span. Then, deriving from that trade-off, we analyze the evolutionary stability of different schedules of age-specific fecundity that are generated by divergent reproductive effort. The analysis was carried out in season lengths that promote either univoltine or bivoltine phenology. The evolutionarily stable reproductive effort decreases with increasing season length in both phenologies, with a sudden increase when a change from univoltine to partially bivoltine phenology takes place. Reproductive effort responds strongly to changing phenology when density-dependent mortality occurs during diapause and weakly when juvenile mortality is density dependent. [ABSTRACT FROM AUTHOR]

Trade-offs have long been a major theme in life-history theory, but they have been hard to document. We introduce a new method that reveals patterns of divergent trade-offs after adjusting for the pervasive variation in rate of resource allocation to offspring as a function of body size and lifestyle. Results suggest that pre-weaning vulnerability to predation has been the major factor determining how female placental mammals allocate production between a few large and many small offspring within a litter and between a few large litters and many small ones within a reproductive season. Artiodactyls, perissodactyls, cetaceans, and pinnipeds, which give birth in the open on land or in the sea, produce a few large offspring, at infrequent intervals, because this increases their chances of escaping predation. Insectivores, fissiped carnivores, lagomorphs, and rodents, whose offspring are protected in burrows or nests, produce large litters of small newborns. Primates, bats, sloths, and anteaters, which carry their young from birth until weaning, produce litters of one or a few offspring because of the need to transport and care for them. [ABSTRACT FROM AUTHOR]