Variation In Life-history Strategies Research Articles

Diversity in habitat use by the East Asian fourfinger threadfin Eleutheronema rhadinum revealed by otolith Sr:Ca and Ba:Ca profiles

Determining the movement and migration patterns of fish between different environments is crucial for understanding the distribution of fish populations and the ecological mechanisms underlying their spatial dynamics. This study is the first to employ otolith Sr:Ca and Ba:Ca profiles to elucidate habitat use by the commercially valuable estuarine species Eleutheronema rhadinum. Age 1+ fish were present in samples from the East China Sea and South China Sea, caught off China in October to December of 2019. Characterization of both otolith element profiles through a quantitative approach suggested migration plasticity for E. rhadinum, especially during the first year. However, the Ba:Ca profile revealed a more distinct pattern of habitat use compared with the Sr:Ca profile, suggesting that the Ba:Ca ratio is more appropriate for life-history reconstruction of this species. Most individuals appeared to be estuary-dependent, with some appearing to have entered fresh water for a short time during their first year of life. After their first winter, individuals tended to move into coastal waters, and most apparently overwintered in the marine environment. Knowledge of the diverse life-history strategies of this valuable species at different life stages should be incorporated into future conservation and management efforts.

Standard metabolic rate does not associate with age-at-maturity genotype in juvenile Atlantic salmon.

Atlantic salmon (Salmo salar) is a species with diverse life‐history strategies, to which the timing of maturation contributes considerably. Recently, the genome region including the gene vgll3 has gained attention as a locus with a large effect on Atlantic salmon maturation timing, and recent studies on the vgll3 locus in salmon have indicated that its effect might be mediated through body condition and accumulation of adipose tissue. However, the cellular and physiological pathways leading from vgll3 genotype to phenotype are still unknown. Standard metabolic rate is a potentially important trait for resource acquisition and assimilation and we hypothesized that this trait, being a proxy for the maintenance energy expenditure of an individual, could be an important link in the pathway from vgll3 genotype to maturation timing phenotype. As a first step to studying links between vgll3 and the metabolic phenotype of Atlantic salmon, we measured the standard metabolic rate of 150 first‐year Atlantic salmon juveniles of both sexes, originating from 14 different families with either late‐maturing or early‐maturing vgll3 genotypes. No significant difference in mass‐adjusted standard metabolic rate was detected between individuals with different vgll3 genotypes, indicating that juvenile salmon of different vgll3 genotypes have similar maintenance energy requirements in the experimental conditions used and that the effects of vgll3 on body condition and maturation are not strongly related to maintenance energy expenditure in either sex at this life stage.

The telomere regulatory gene POT1 responds to stress and predicts performance in nature: Implications for telomeres and life history evolution.

Telomeres are emerging as correlates of fitness-related traits and may be important mediators of ecologically relevant variation in life history strategies. Growing evidence suggests that telomere dynamics can be more predictive of performance than length itself, but very little work considers how telomere regulatory mechanisms respond to environmental challenges or influence performance in nature. Here, we combine observational and experimental data sets from free-living tree swallows (Tachycinetabicolor) to assess how performance is predicted by the telomere regulatory gene POT1, which encodes a shelterin protein that sterically blocks telomerase from repairing the telomere. First, we show that lower POT1 gene expression in the blood was associated with higher female quality, that is, earlier breeding and heavier body mass. We next challenged mothers with an immune stressor (lipopolysaccharide injection) that led to "sickness" in mothers and 24h of food restriction in their offspring. While POT1 did not respond to maternal injection, females with lower constitutive POT1 gene expression were better able to maintain feeding rates following treatment. Maternal injection also generated a 1-day stressor for chicks, which responded with lower POT1 gene expression and elongated telomeres. Other putatively stress-responsive mechanisms (i.e., glucocorticoids, antioxidants) showed marginal responses in stress-exposed chicks. Model comparisons indicated that POT1 mRNA abundance was a largely better predictor of performance than telomere dynamics, indicating that telomere regulators may be powerful modulators of variation in life history strategies.

Interpopulation differences in male reproductive effort drive the population dynamics of a host exposed to an emerging fungal pathogen.

Compensatory recruitment is a key demographic mechanism that has allowed the coexistence of populations of susceptible amphibians with Batrachochytrium dendrobatidis (Bd), a fungus causing one of the most devastating emerging infectious disease ever recorded among vertebrates. However, the underlying processes (e.g. density-dependent increase in survival at early life stages, change in reproductive traits) as well as the level of interpopulation variation in this response are poorly known. We explore potential mechanisms of compensatory recruitment in response to Bd infection by taking advantage of an amphibian system where male reproductive traits are easy to quantify in free-living populations. The Southern Darwin's frog Rhinoderma darwinii is a vocal sac-brooding species that exhibits a high susceptibility to lethal Bd infection. Using a 7-year capture-recapture study at four populations with contrasting Bd infection status (one high prevalence, one low prevalence and two Bd-free populations), we evaluated whether Bd-positive populations exhibited a higher adult recruitment and a higher male reproductive effort than Bd-negative populations. We also estimated population growth rates to explore whether recruitment compensated for the negative impacts of Bd on the survival of adults. In addition, we evaluated a potential demographic signal of compensatory recruitment (i.e. positive relationship between the proportion of juveniles and Bd prevalence) in response to Bd infection using raw count data from 13 R. darwinii populations. The high Bd prevalence population exhibited the highest male reproductive effort and the highest recruitment among the four monitored populations. This led to a growing population during the study period despite high mortality of adult hosts. In contrast, males from the population with low Bd prevalence had a low reproductive effort and this population, which had the lowest adult recruitment, was declining during the study period despite adults having a higher survival in comparison to the high Bd prevalence population. We also found a demographic signal of compensatory recruitment in response to Bd infection in our broader analysis of 13 R. darwinii populations. Our study underlines the importance of interpopulation variation in life-history strategies on the fate of host populations after infectious disease emergence. Our results also suggest that an increase in reproductive effort can be one of the processes underlying compensatory recruitment in populations of Bd-susceptible amphibians.

An empirical confirmation of diversified bet hedging as a survival strategy in unpredictably varying environments.

Environmental change jeopardizes the survival of species from variable environments by making the occurrence of favorable conditions less predictable. For organisms with long-lived propagules (e.g., spores, eggs, or seeds), the theory of diversified bet hedging (DBH) predicts that delayed hatching over different growing seasons can help populations avoid extinction. Empirical observations in different organisms are consistent with DBH, but integrated tests that simultaneously validate the main theoretical assumptions and predictions are lacking. In this study, we combine field and multi-generational lab experiments to provide a complete test of DBH. Consistent with DBH predictions, resting egg clutches of the fairy shrimp Branchipodopsis wolfi, which inhabits rain-fed temporary rock pool environments with unpredictable inundations, hatched partially over a succession of inundations with identical hatching cues. Bet hedging was more common in populations from more unpredictable habitats where hatching fractions were lower. This differentiation in hatching strategies was preserved after two generations under common garden conditions, which implies intrinsic (epi-)genetic control of hatching. Finally, a demographic model confirmed that lower hatching fractions increase long-term population growth in unpredictable habitats. With this paper we propose a method to calculate probabilities of successful recruitment for organisms that use imperfect cues and show that this drives selection for variation in life history strategies as part of a DBH strategy.

Early-life seasonal, weather and social effects on telomere length in a wild mammal.

Early-life environmental conditions can provide a source of individual variation in life-history strategies and senescence patterns. Conditions experienced in early life can be quantified by measuring telomere length, which can act as a biomarker of survival probability in some species. Here, we investigate whether seasonal changes, weather conditions and group size are associated with early-life and/or early-adulthood telomere length in a wild population of European badgers (Meles meles). We found substantial intra-annual changes in telomere length during the first 3years of life, where within-individual effects showed shorter telomere lengths in the winter following the first spring and a trend for longer telomere lengths in the second spring compared to the first winter. In terms of weather conditions, cubs born in warmer, wetter springs with low rainfall variability had longer early-life (3-12months old) telomeres. Additionally, cubs born in groups with more cubs had marginally longer early-life telomeres, providing no evidence of resource constraint from cub competition. We also found that the positive association between early-life telomere length and cub survival probability remained when social and weather variables were included. Finally, after sexual maturity, in early adulthood (i.e., 12-36months) we found no significant association between same-sex adult group size and telomere length (i.e., no effect of intrasexual competition). Overall, we show that controlling for seasonal effects, which are linked to food availability, is important in telomere length analyses, and that variation in telomere length in badgers reflects early-life conditions and also predicts first year cub survival.

Morph-specific life-history correlations in a wing-dimorphic water strider.

Life-history theory predicts a negative correlation between reproduction and survival because individuals differ in their investment in early reproduction at the expense of survival. However, life-history trade-offs can be masked when individual differences in resource allocation are smaller than those in resource acquisition. In polymorphic species, as distinct morphs exhibit differences in intrinsic physiology, the relative effects of resource acquisition and allocation on life-history traits will differ between morphs, contributing to morph-specific life-history correlations. Here, in the wing-dimorphic water strider Aquarius paludum, we found that wing morphs differed in within-morph individual-level life-history correlations. Longer-lived flight-capable long-winged females produced fewer eggs per day and matured later, whereas life-history trade-offs were not observed in short-winged flightless females. The survival-reproduction trade-off observed in long-winged females may be a result of individual differences in the timing of wing muscle histolysis. Individuals that underwent wing histolysis early would have increased reproduction at the expense of a shorter life, whereas individuals with late wing histolysis would have reduced reproduction but a longer life. Short-winged females, who never develop wings, effectively have more resources to allocate to both survival and reproduction, masking any life-history trade-offs. Thus, we suggest that morph-specific effects of resource allocation trade-offs can shape the morph-specific extent of individual variation in life-history strategies, which may contribute to the evolution and maintenance of within-species polymorphism.

Microbial dysbiosis reflects disease resistance in diverse coral species

Disease outbreaks have caused significant declines of keystone coral species. While forecasting disease outbreaks based on environmental factors has progressed, we still lack a comparative understanding of susceptibility among coral species that would help predict disease impacts on coral communities. The present study compared the phenotypic and microbial responses of seven Caribbean coral species with diverse life-history strategies after exposure to white plague disease. Disease incidence and lesion progression rates were evaluated over a seven-day exposure. Coral microbiomes were sampled after lesion appearance or at the end of the experiment if no disease signs appeared. A spectrum of disease susceptibility was observed among the coral species that corresponded to microbial dysbiosis. This dysbiosis promotes greater disease susceptiblity in coral perhaps through different tolerant thresholds for change in the microbiome. The different disease susceptibility can affect coral’s ecological function and ultimately shape reef ecosystems.

Viral load, not food availability or temperature, predicts colony longevity in an invasive eusocial wasp with plastic life history

Social insect colonies exhibit a variety of life history strategies, from the annual, semelparous colonies of temperate bees and wasps to the long-lived colonies of many ants and honeybees. Species introduced to novel habitats may exhibit plasticity in life history strategies as a result of the introduction, but the factors governing these changes often remain obscure. Vespula pensylvanica, a yellowjacket wasp, exhibits such plasticity in colony longevity. Multi-year (perennial) colonies are relatively common in introduced populations in Hawaii, while source populations in the western United States are typically on an annual cycle. Here, we use experiments and observational data to examine how diet, disease, nest thermal environment, and nest location influence colony longevity in a population with both annual and perennial colonies. Counter to our predictions, experimental feeding and warming did not increase colony survival in the winter in the introduced range. However, Moku Virus load and wasp colony density predicted colony survival in one year, suggesting a potential role for disease in modulating colony phenology. We also found that local V. pensylvanica colony density was positively correlated with Moku Virus loads, and that Arsenophonus sp. bacterial loads in V. pensylvanica colonies were positively associated with proximity to feral honeybee (Apis mellifera) hives, suggesting potential transmission routes for these poorly understood symbionts. The factors influencing colony longevity in this population are likely multiple and interactive. More important than food availability, we propose winter precipitation as a critical factor that may explain temporal and spatial variation in colony longevity in these invasive wasps.

Nest microclimate and limits to egg viability explain avian life-history variation across latitudinal gradients.

Variation in life-history strategies is central to our understanding of population dynamics and how organisms adapt to their environments. Yet we lack consensus regarding the ecological processes that drive variation in traits related to reproduction and survival. For example, we still do not understand the cause of two widespread inter- and intraspecific patterns: (1) the ubiquitous positive association between avian clutch size and latitude; and (2) variation in the extent of asynchronous hatching of eggs within a single clutch. Well-known hypotheses to explain each pattern have largely focused on biotic processes related to food availability and predation risk. However, local adaptation to maintain egg viability could explain both patterns with a single abiotic mechanism. The egg viability hypothesis was initially proposed to explain the cause of asynchronous hatching and suggests that asynchronous hatching results from early incubation onset in response to unfavorable nest microclimatic conditions, which otherwise reduce egg viability. However, allocation of resources to early incubation, prior to clutch completion, may energetically constrain clutch size and help explain the positive association between clutch size and latitude. We measured intraspecific variation in five functionally linked life-history traits of burrowing owls at five study sites spanning a 1,400-km latitudinal transect in western North America: clutch size, the timing of incubation onset, the degree of hatching asynchrony, the probability of hatching failure, and nestling survival. We found that most traits varied clinally with latitude, but all the traits were more strongly associated with individual nest microclimates than with latitude, and all varied with nest microclimate in the directions predicted by the egg viability hypothesis. Furthermore, incubation began earlier, hatching asynchrony increased, and clutch size declined across the breeding season. These results suggest that nest microclimate drives an important life-history trade-off and that thermal gradients are often sufficient to account for observed biogeographic and seasonal patterns in life-history strategies. Furthermore, our results reveal a potentially important indirect mechanism by which reproductive success and recruitment could be affected by climate change.

Progress on Research Regarding Ecology and Biodiversity of Coastal Fisheries and Nektonic Species and Their Habitats within Coastal Landscapes

This paper aims to highlight the new research and significant advances in our understanding of links between coastal habitat quality/quantity/diversity and the diversity of fisheries species and other mobile aquatic species (hereafter nekton) that use them within coastal landscapes. This topic is quite diverse owing to the myriad of habitat types found in coastal marine waters and the variety of life history strategies fisheries species and nekton use in these environments. Thus, we focus our review on five selective but relevant topics, habitat templates, essential fish habitat, habitat mosaics/habitat connectivity, transitory/ephemeral habitat, and the emerging/maturing approaches to the study of fish-habitat systems as a roadmap to its development. We have highlighted selected important contributions in the progress made on each topic to better identify and quantify landscape scale interactions between living biota and structured habitats set within a dynamic landscape.

Differing Life-History Strategies of Two Mycoheterotrophic Orchid Species Associated with Leaf Litter- and Wood-Decaying Fungi

Mycoheterotrophic orchids depend completely on mycorrhizal fungi for their supply of carbon. The life-history traits of mycoheterotrophic plants (MHPs) can differ according to the characteristics of the associated mycorrhizal fungi. We compared the life-history strategies of two mycoheterotrophic orchids associated with wood- and leaf litter-decaying fungi over a maximum of six years of field monitoring. Seventy percent of the aboveground stems of Erythrorchis altissima, associated with wood-decaying fungi, disappeared from the host wood within two years after tagging, likely due to nutrient depletion. In contrast, Gastrodia confusa, associated with leaf litter-decaying fungi, occurred continuously (18 to 108 fruiting stalks) every year within a small-scale plot (12 × 45 m) for six years through seed and clonal propagation. Our results support the idea that mycoheterotrophic orchids associated with wood-decaying fungi disappear from their habitats due to nutrient depletion after their host wood has mostly decayed, while mycoheterotrophic orchids associated with leaf litter-decaying fungi can survive in small-scale habitats where substantial leaf fall regularly occurs to sustain the associated fungi. Our study provides basic information about a unique life-history strategy in MHPs associated with saprotrophic fungi and an understanding of the variation in life-history strategies among MHPs.

Nest-Site Choice and Breeding Success among Four Sympatric Species of Passerine Birds in a Reedbed-Dominated Wetland

The efficacy of nest-site choice ultimately determines the breeding success of birds. Comparisons of the reproductive strategies of various bird species which inhabit the same habitat may provide insights on the evolution of the diverse life-history strategies in birds. In this study, nest-site choice and breeding success of four species of passerine birds that rely on reeds for reproduction were investigated in a wetland in Hebei, China. The four species were the Oriental reed warbler (Acrocephalus orientalis) (ORW), the blunt-winged warbler (Acrocephalus concinens) (BW), the reed parrotbill (Paradoxornis heudei) (RP), and the vinous-throated parrotbill (Sinosuthora webbiana) (VP). Our results showed that breeding nests of the four species were distributed in a mosaic pattern within the same habitat, with similar nest shapes/structures and nest-sites in the reeds. The only characteristics which differed significantly among the species were nest height above the water surface, height of reeds where nests were placed, and density of reeds around the nest site. In addition, the starting time of reproduction clearly differed for the four species. The breeding success rates of the four species were 34.5% (86/249) for ORW, 35.3% (6/17) for BW, 38.5% (15/39) for RP, and 40.9% (9/22) for VP in the two study years. The main factors affecting the breeding success were nest predation and poor weather conditions, like heavy rainstorms and wind, while nest parasitism by the common cuckoo (Cuculus canorus) represented an important factor for breeding failure only in the Oriental reed warbler. Our study demonstrated that these four sympatric species of passerine birds inhabiting the same wetland exhibit differences in terms of nest-site choice and breeding phenology.

Cis-regulatory differences in isoform expression associate with life history strategy variation in Atlantic salmon.

A major goal in biology is to understand how evolution shapes variation in individual life histories. Genome-wide association studies have been successful in uncovering genome regions linked with traits underlying life history variation in a range of species. However, lack of functional studies of the discovered genotype-phenotype associations severely restrains our understanding how alternative life history traits evolved and are mediated at the molecular level. Here, we report a cis-regulatory mechanism whereby expression of alternative isoforms of the transcription co-factor vestigial-like 3 (vgll3) associate with variation in a key life history trait, age at maturity, in Atlantic salmon (Salmo salar). Using a common-garden experiment, we first show that vgll3 genotype associates with puberty timing in one-year-old salmon males. By way of temporal sampling of vgll3 expression in ten tissues across the first year of salmon development, we identify a pubertal transition in vgll3 expression where maturation coincided with a 66% reduction in testicular vgll3 expression. The late maturation allele was not only associated with a tendency to delay puberty, but also with expression of a rare transcript isoform of vgll3 pre-puberty. By comparing absolute vgll3 mRNA copies in heterozygotes we show that the expression difference between the early and late maturity alleles is largely cis-regulatory. We propose a model whereby expression of a rare isoform from the late allele shifts the liability of its carriers towards delaying puberty. These results exemplify the potential importance of regulatory differences as a mechanism for the evolution of life history traits.

Geographic Variation in Life-History Traits of Black Sea Bass (Centropristis striata) During a Rapid Range Expansion

The warming of the world’s oceans has resulted in the redistribution of many marine species globally. As species undergo range shifts, the expanding edge of the population often experiences novel environmental and demographic conditions that may result in the emergence of variation in life-history strategies. The northern stock of black sea bass, Centropristis striata, has recently expanded its distribution poleward, into the Gulf of Maine. Management has struggled to keep pace with this rapid range shift, in part, because very little is known about the expanding population. We compared life-history traits of black sea bass collected from 2013-2016 from the northern most point of the historic range of the northern stock (southern Massachusetts) to those from two areas in the newly expanded range (northern Massachusetts and Maine). We found significant differences in size, diet, condition, maturity and sex ratio between black sea bass from southern Massachusetts and the Gulf of Maine. Overall, sea bass in the newly expanded range consumed a less diverse diet and their condition was lower, but they reached maturity at a younger age. We also found greater length- and age-at-maturity estimates from all regions combined compared to the most recent black sea bass stock assessment which includes data from Cape Hatteras, NC to southern Massachusetts. This study represents initial observations of life-history traits of sea bass in its newly expanded range in the Gulf of Maine, and suggests that these sea bass exhibit life-history strategies that differ from their southern counterparts within their historic range. Given these findings, the stock assessment for the Northeast U.S. Continental Shelf black sea bass stock may not be adequate for sea bass in the Gulf of Maine. Studies investigating the expanding edge of economically valuable fishery species are needed to aid in ongoing and future efforts to assess and manage their stocks.

An ecological driver for the macroevolution of morphological polymorphism within colonial invertebrates.

Colonial marine invertebrates, such as corals and bryozoans, have modular growth. Individual modules within a colony are homologous to an individual solitary animal body. But in contrast to the predominately sexual origin of solitary animal bodies, modules within a colony are always produced asexually. The repetition of modules and the indeterminism of their organization gives colonies the ability to grow in ways solitary animals cannot. Colonial invertebrates consequently grow in such a way as to resemble weeds, bushes, or trees. The multitude of growth forms of colonial invertebrates arise from differences how individual colonies within a species tend to invest their energy into modular growth, persistence, asexual propagation, and sexual reproduction. Moreover, many colonial invertebrates possess several body types, morphological polymorphism among modules, where modules qualitatively differ in shape, size, and function. In this paper, I propose a mechanism that links the origin of novel body types to the evolution of life‐history strategies among species. When colonies first evolve from solitary ancestors, the life‐history strategy of the colony remains constrained by the life‐history strategies of the individual modules within the colony until a new polymorph type evolves. The addition of novel body types within a colony introduces potential variation in life‐history strategies. Colonies can then change strategies by regulating the frequencies of body types within the colony. This, along with the ability of body types to simplify their structure permits colonies to evolve more complex life‐histories. Each new polymorph type that evolves permits more variation in colonial life‐histories to exist.

Child mortality and parental grief: An evolutionary analysis

Across most of human history, infant and child mortality rates were very high, suggesting the death of a child was a challenge faced by many ancestral parents. Prolonged grief likely harmed grievers’ fitness, yet grief is ubiquitous and often protracted, thereby presenting a puzzle for evolutionary arguments. We integrate existing theories of grief with patterns of parental bereavement to examine how human psychology has been shaped to respond to the death of a child. We contend that variation in life history strategy may explain the relative difficulty with which individuals recover from losing a child. We propose that the same physiological mechanisms underlying detachment and grief during dissolved romantic relationships may also underlie the intensity of parental attachment and bereavement. This theoretical review thus integrates evolutionary theory with extant grief research to provide a functional analysis of the immense suffering associated with the loss of a child.

Life history evolution, species differences, and phenotypic plasticity in hemiparasitic eyebrights (Euphrasia).

Species delimitation in parasitic organisms is challenging because traits used to identify species are often plastic and vary depending on the host. Here, we use species from a recent radiation of generalist hemiparasitic Euphrasia to investigate trait variation and trait plasticity. We tested whether Euphrasia species show reliable trait differences, investigated whether these differences correspond to life history trade-offs between growth and reproduction, and quantified plasticity in response to host species. Common garden experiments were used to evaluate trait differences between 11 Euphrasia taxa grown on a common host, document phenotypic plasticity when a single Euphrasia species is grown on eight different hosts, and relate observations to trait differences recorded in the wild. Euphrasia exhibited variation in life history strategies; some individuals transitioned rapidly to flowering at the expense of early season growth, while others invested in vegetative growth and delayed flowering. Life history differences were present between some species, though many related taxa lacked clear trait differences. Species differences were further blurred by phenotypic plasticity-many traits were plastic and changed with host type or between environments. Phenotypic plasticity in response to host and environment confounds species delimitation in Euphrasia. When grown in a common garden environment, some morphologically distinct taxa can be identified, though others represent morphologically similar shallow segregates. Trait differences present between some species and populations demonstrate the rapid evolution of distinct life history strategies in response to local ecological conditions.

RETRACTED ARTICLE: Novel life history strategy in a deep sea fish challengesassumptions about reproduction in extreme environments

The deep ocean is frequently assumed to be a homogeneous system lackingthe same diverse life history strategies found in shallower waters. However, as ourmethods for exploring the deep ocean improve, common assumptions about dispersal,reproduction and behavior are constantly being challenged. Fishes exhibit the mostdiverse reproductive strategies among vertebrates. Understanding life historystrategies in deep-sea environments is lacking for many species of fishes. Here, wereport a novel reproductive strategy where a fish (Parazenpacificus) provides parental care via mouth brooding. This behavioris observed from a specimen collected with eggs present in the buccal cavity, alongwith other specimens exhibiting pre-brooding morphologies. This is the firstdescription of this unique life history trait in a deep-sea fish and fills in a gapin the larval literature for this family of fishes and prompts further investigationinto other novel reproductive modes of deep-sea fauna.

Life‐history strategy varies with the strength of competition in a food‐limited ungulate population

Fluctuating population density in stochastic environments can contribute to maintain life-history variation within populations via density-dependent selection. We used individual-based data from a population of Soay sheep to examine variation in life-history strategies at high and low population density. We incorporated life-history trade-offs among survival, reproduction and body mass growth into structured population models and found support for the prediction that different life-history strategies are optimal at low and high population densities. Shorter generation times and lower asymptotic body mass were selected for in high-density environments even though heavier individuals had higher probabilities to survive and reproduce. In contrast, greater asymptotic body mass and longer generation times were optimal at low population density. If populations fluctuate between high density when resources are scarce, and low densities when they are abundant, the variation in density will generate fluctuating selection for different life-history strategies, that could act to maintain life-history variation.