Growth-mortality Trade-off Research Articles

Leaf and twig traits predict habitat adaptation and demographic strategies in tropical freshwater swamp forest trees.

Differences in demographic and environmental niches facilitate plant species coexistence in tropical forests. However, the adaptations that enable species to achieve higher demographic rates (e.g. growth or survival) or occupy unique environmental niches (e.g. waterlogged conditions) remain poorly understood. Anatomical traits may better predict plant environmental and demographic strategies because they are direct measurements of structures involved in these adaptations. We collected 18 leaf and twig traits from 29 tree species in a tropical freshwater swamp forest in Singapore. We estimated demographic parameters of the 29 species from growth and survival models, and degree of association toward swamp habitats. We examined pairwise trait-trait, trait-demography and trait-environment links while controlling for phylogeny. Leaf and twig anatomical traits were better predictors of all demographic parameters than other commonly measured leaf and wood traits. Plants with wider vessels had faster growth rates but lower survival rates. Leaf and spongy mesophyll thickness predicted swamp association. These findings demonstrate the utility of anatomical traits as indicators of plant hydraulic strategies and their links to growth-mortality trade-offs and waterlogging stress tolerance that underlie species coexistence mechanisms in tropical forest trees.

Basin-wide variation in tree hydraulic safety margins predicts the carbon balance of Amazon forests

Tropical forests face increasing climate risk1,2, yet our ability to predict their response to climate change is limited by poor understanding of their resistance to water stress. Although xylem embolism resistance thresholds (for example, varPsi 50) and hydraulic safety margins (for example, HSM50) are important predictors of drought-induced mortality risk3–5, little is known about how these vary across Earth’s largest tropical forest. Here, we present a pan-Amazon, fully standardized hydraulic traits dataset and use it to assess regional variation in drought sensitivity and hydraulic trait ability to predict species distributions and long-term forest biomass accumulation. Parameters varPsi 50 and HSM50 vary markedly across the Amazon and are related to average long-term rainfall characteristics. Both varPsi 50 and HSM50 influence the biogeographical distribution of Amazon tree species. However, HSM50 was the only significant predictor of observed decadal-scale changes in forest biomass. Old-growth forests with wide HSM50 are gaining more biomass than are low HSM50 forests. We propose that this may be associated with a growth–mortality trade-off whereby trees in forests consisting of fast-growing species take greater hydraulic risks and face greater mortality risk. Moreover, in regions of more pronounced climatic change, we find evidence that forests are losing biomass, suggesting that species in these regions may be operating beyond their hydraulic limits. Continued climate change is likely to further reduce HSM50 in the Amazon6,7, with strong implications for the Amazon carbon sink.

Predicting ecosystem productivity based on plant community traits.

With the rapid accumulation of plant trait data, major opportunities have arisen for the integration of these data into predicting ecosystem primary productivity across a range of spatial extents. Traditionally, traits have been used to explain physiological productivity at cell, organ, or plant scales, but scaling up to the ecosystem scale has remained challenging. Here, we show the need to combine measures of community-level traits and environmental factors to predict ecosystem productivity at landscape or biogeographic scales. We show how theory can extend the production ecology equation to enormous potential for integrating traits into ecological models that estimate productivity-related ecosystem functions across ecological scales and to anticipate the response of terrestrial ecosystems to global change.

Ecological mechanisms generating variable first-year growth in Paralichthys lethostigma, a coastal flatfish

Determining the relative influence of, and possible synergistic relationships among, environmental and biotic processes on individual growth rates can identify key mechanisms that contribute to variation in body size observed among similar-aged individuals. The timing of ontogenetic shifts in diet or habitat, which themselves can be size-dependent, can have considerable impacts on individual growth trajectories, survival, and year class formation. The southern flounder Paralichthys lethostigma is a coastal flatfish species with juvenile cohorts that occupy diverse estuarine settlement habitats and commonly display 2-3 fold variation in body size by the end of the first growing season. We examined abiotic and biotic factors contributing to differences in early juvenile growth for the 2018 year-class of southern flounder in a North Carolina estuary. Following peak larval settlement (Febuary/March), flounder body sizes began to diverge by April, with differences between settlement habitats contributing to roughly 3-fold variation by August. Stomach content analyses indicated a greater occurrence of fish prey in the diets of southern flounder recovered in mesohaline versus oligohaline settlement habitats. Increased piscivorous foraging in mesohaline habitats was linked to nitrogen (δ15N)-enriched tissues and faster growth rates relative to fish collected in oligohaline habitats. Results from a controlled laboratory experiment provided evidence for the strong effects of diet, with only minor direct effects of salinity on juvenile southern flounder growth. Our findings support the critical role of ontogenetic shifts to piscivorous feeding for growth in young-of-the-year southern flounder and highlights the potential for growth-mortality tradeoffs to exist among estuarine settlement habitats.

The interspecific growth-mortality trade-off is not a general framework for tropical forest community structure.

Resource allocation within trees is a zero-sum game. Unavoidable trade-offs dictate that allocation to growth-promoting functions curtails other functions, generating a gradient of investment in growth versus survival along which tree species align, known as the interspecific growth-mortality trade-off. This paradigm is widely accepted but not well established. Using demographic data for 1,111 tree species across ten tropical forests, we tested the generality of the growth-mortality trade-off and evaluated its underlying drivers using two species-specific parameters describing resource allocation strategies: tolerance of resource limitation and responsiveness of allocation to resource access. Globally, a canonical growth-mortality trade-off emerged, but the trade-off was strongly observed only in less disturbance-prone forests, which contained diverse resource allocation strategies. Only half of disturbance-prone forests, which lacked tolerant species, exhibited the trade-off. Supported by a theoretical model, our findings raise questions about whether the growth-mortality trade-off is a universally applicable organizing framework for understanding tropical forest community structure.

Personality-driven life history trade-offs differ in two subpopulations of free-ranging predators.

Consistent individual differences in behaviour (i.e. personality) can be explained in an evolutionary context if they are favoured by life history trade-offs as conceptualized in the pace-of-life syndrome (POLS) hypothesis. Theory predicts that faster-growing individuals suffer higher mortality and that this trade-off is mediated through exploration/risk-taking personality, but empirical support for this remains limited and ambiguous. Equivocal support to the POLS hypothesis suggests that the link between life history and personality may only emerge under certain circumstances. Understanding personality-driven trade-offs would be facilitated by long-term studies in wild populations experiencing different ecological conditions. Here, we tested whether personality measured in semi-captivity was associated with a growth-mortality trade-off via risk-taking in the wild in two subpopulations of juvenile lemon sharks Negaprion brevirostris known to differ in their predator abundance. We expected stronger personality-driven trade-offs in the predator-rich environment as compared to the predator-poor environment. Sharks were captured yearly from 1995 onwards allowing us to obtain long-term data on growth and apparent survival in each subpopulation. We then used a novel open-field assay to test sharks for exploration personality yearly from 2012 to 2017. A subset of the tested sharks was monitored in the field using telemetry to document risk-taking behaviours. We tested (a) if fast explorers in captivity took more risks and grew faster in the wild and (b) if natural selection acted against more explorative, faster-growing sharks. In the subpopulation with fewer predators, more explorative sharks in captivity took more risks in the wild and grew faster. In turn, larger, fast-growing sharks had lower apparent survival. In the predator-rich subpopulation, despite finding selection on fast growth, we found no link between exploration personality and the growth-mortality trade-off. Our study demonstrates that the association between personality and life history is favoured in some ecological contexts but not in others. We identify predator and resource abundance as two main potential drivers of the personality-mediated trade-off and emphasize that future work on the POLS hypothesis would benefit from an approach integrating behaviour and life history across ecological conditions.

The geometry of reaction norms yields insights on classical fitness functions for Great Lakes salmon.

Life history theory examines how characteristics of organisms, such as age and size at maturity, may vary through natural selection as evolutionary responses that optimize fitness. Here we ask how predictions of age and size at maturity differ for the three classical fitness functions–intrinsic rate of natural increase r, net reproductive rate R0, and reproductive value Vx−for semelparous species. We show that different choices of fitness functions can lead to very different predictions of species behavior. In one’s efforts to understand an organism’s behavior and to develop effective conservation and management policies, the choice of fitness function matters. The central ingredient of our approach is the maturation reaction norm (MRN), which describes how optimal age and size at maturation vary with growth rate or mortality rate. We develop a practical geometric construction of MRNs that allows us to include different growth functions (linear growth and nonlinear von Bertalanffy growth in length) and develop two-dimensional MRNs useful for quantifying growth-mortality trade-offs. We relate our approach to Beverton-Holt life history invariants and to the Stearns-Koella categorization of MRNs. We conclude with a detailed discussion of life history parameters for Great Lakes Chinook Salmon and demonstrate that age and size at maturity are consistent with predictions using R0 (but not r or Vx) as the underlying fitness function.

Shade tolerance within the context of the successional process in tropical rain forests

Shade tolerance (the capacity to survive and grow over long periods under shade) is a key component of plant fitness and the foundation of current theories of forest succession in tropical rain forests. It serves as a paradigm to understand the optimal allocation of limited resources under dynamic light regimes. I analyze how tropical rain forest succession influences the expression of ecophysiological mechanisms leading to shade tolerance, and identify future areas that will increase our understanding of the ecological and evolutionary consequences of this phenomenon. Shade tolerance is a multivariate, continuous functional trait reflecting the growth-mortality trade-off of investing resources under limited light vs. exploiting high light conditions. I propose the life cycle successional trajectory model of Gómez-Pompa & Vázquez-Yanes as an integrative tool to understand tropical rain forest succession. This model shows how species distribute along the successional environmental gradient based on their degree of shade tolerance and represents a more integrative paradigm to understand the interface between different aspects of species diversity (ontogenetic variation and functional diversity) throughout succession. It proposes that different trait combinations determining shade tolerance are expressed at different stages of the life cycle, which affects how and when plants enter the successional trajectory. Models explaining the expression of shade tolerance (resource availability, carbon gain, CSR, resource competition) are based on whole-plant economics and are not mutually exclusive. The analysis of shade tolerance is biased towards tree seedlings in the understory of mature forests. Other life stages (juvenile and adult trees), life forms, and microhabitats throughout the forest profile are almost always excluded from these analyses. More integrative explanations based on the distribution of functional traits among species, ontogenetic stages, and the nature of the environmental gradient are being developed based on long-term data and chronosequence comparisons. In summary, shade-tolerance is a complex phenomenon, is determined by multiple characters that change ontogenetically over space and time and entails considerable plasticity. Current methods do not account for this plasticity. Understanding the nature of shade tolerance and its functional basis is critical to comprehending plant performance and improving the management, restoration and conservation of tropical rain forests given the combined threats of global warming and habitat loss

Density-dependent survival varies with species life-history strategy in a tropical forest.

Species coexistence in diverse communities likely results from multiple interacting factors. Mechanisms such as conspecific negative density dependence (CNDD) and varying life-history strategies related to resource partitioning are known to influence plant fitness, and thereby community composition and diversity. However, we have little understanding of how these mechanisms interact and how they vary across life stages. Here, we document the interaction between CNDD and life-history strategy, based on growth-mortality trade-offs, from seedling to adult tree for 47 species in a tropical forest. Species' life-history strategies remained consistent across stages: fast-growing species had higher mortality than slow-growing species at all stages. In contrast, mean CNDD was strongest at early life stages (i.e. seedling, sapling). Fast-growing species tended to suffer greater CNDD than slow-growing species at several, but not all life stages. Overall, our results demonstrate that coexistence mechanisms interact across multiple life stages to shape diverse tree communities.

Xylem hydraulic safety and construction costs determine tropical tree growth.

Faster growth in tropical trees is usually associated with higher mortality rates, but the mechanisms underlying this relationship are poorly understood. In this study, we investigate how tree growth patterns are linked with environmental conditions and hydraulic traits, by monitoring the cambial growth of 9 tropical cloud forest tree species coupled with numerical simulations using an optimization model. We find that fast-growing trees have lower xylem safety margins than slow-growing trees and this pattern is not necessarily linked to differences in stomatal behaviour or environmental conditions when growth occurs. Instead, fast-growing trees have xylem vessels that are more vulnerable to cavitation and lower density wood. We proposethe growth - xylem vulnerability trade-off represents a wood hydraulic economics spectrum similar to the classic leaf economic spectrum, and show through numerical simulations that this trade-off can emerge from the coordination between growth rates, wood density, and xylem vulnerability to cavitation. Our results suggest that vulnerability to hydraulic failure might be related with the growth-mortality trade-off in tropical trees, determining important life history differences. These findings are important in furthering our understanding of xylem hydraulic functioning and its implications on plant carbon economy.

Sea trout adapt their migratory behaviour in response to high salmon lice concentrations.

Sea trout face growth-mortality trade-offs when entering the sea to feed. Salmon lice epizootics resulting from aquaculture have shifted these trade-offs, as salmon lice might both increase mortality and reduce growth of sea trout. We studied mortality and behavioural adaptations of wild sea trout in a large-scale experiment with acoustic telemetry in an aquaculture intensive area that was fallowed (emptied of fish) synchronically biannually, creating large variations in salmon lice concentrations. We tagged 310 wild sea trout during 3years, and gave half of the individuals a prophylaxis against further salmon lice infestation. There was no difference in survival among years or between treatments. In years of high infestation pressure, however, sea trout remained closer to the river outlet, used freshwater (FW) habitats for longer periods and returned earlier to the river than in the low infestation year. This indicates that sea trout adapt their migratory behaviour by actively choosing FW refuges from salmon lice to escape from immediate mortality risk. Nevertheless, simulations show that these adaptations can lead to lost growth opportunities. Reduced growth can increase long-term mortality of sea trout due to prolonged exposure to size-dependent predation risk, lead to lower fecundity and, ultimately, reduce the likelihood of sea migration.

Biotic and abiotic drivers of the tree growth and mortality trade-off in an old-growth temperate forest

The tree growth and mortality trade-off is well documented and plays a key role in forests by forming the basis of many ecosystem processes and contributing to tree species coexistence. One area that requires further understanding is how the growth-mortality trade-off is influenced by biotic and abiotic interactions in the forest. In this study, we used a 5-year interval of tree growth and mortality demographic data from a 9-ha forest dynamics plot in a temperate old-growth Chinese forest to address two questions, (1) What is the nature of the growth-mortality trade-off among species in this forest? (2) Are there differences between the responses of tree growth and mortality to local neighborhood variables, both biotic and abiotic? Specifically, do these responses vary among species and with tree size? For the first question, we hypothesized that, within species, mortality rate would be negatively correlated with growth rate, whereas among species it would be positively correlated. For the second question, we expected biotic and abiotic factors to both be important for tree mortality; but expected biotic factors to be more important than abiotic factors for tree growth. The responses of tree growth and mortality to local neighborhood variables, both biotic and abiotic, varied among species. Abiotic factors were more important than biotic factors for small trees; biotic variables were stable with tree size for both small and large trees. Our results showed that there were differences between the responses of tree growth and mortality to local neighborhood variables (i.e., biotic and abiotic) in this temperate forest community. The responses of tree mortality to local neighborhood variables strongly varied across species. Abiotic factors were only important for the growth of small trees. Biotic factors were stable with tree size for growth of small and large trees, but not for mortality. Mortality was negatively correlated with growth for large trees within species; growth-mortality trade-off among species was found to be more accentuated for small trees. The responses of tree mortality to local neighborhood variables differed strongly across species, whereas the responses of tree growth to local neighborhood variables varied among size classes. Tree growth and mortality is tightly correlated both within (negatively) and among species (positively), and their relationships are both size dependent in this temperate forest. Our findings highlight that differential responses of tree growth and mortality to local neighborhood variables continue to shape the forest community well after sapling establishment in this temperate forest community.

Inactive trout come out at night: behavioral variation, circadian activity, and fitness in the wild.

Theory suggests that high activity levels in animals increase growth at the cost of increased mortality. This growth-mortality tradeoff has recently been incorporated into the wider framework of the pace-of-life syndrome (POLS) hypothesis. However, activity is often quantified only in the laboratory and on a diurnal basis, leaving open the possibility that animals manage predation risk and feeding efficiency in the wild by modulating their circadian activity rhythms. Here we investigate how laboratory activity in wild brown trout parr (Salmo trutta L.) associates with circadian activity, growth, and mortality in their natal stream. We found that individuals with high activity in the laboratory displayed high dispersal and cathemeral activity in their natal stream. In contrast, trout with low laboratory activity showed variation of activity in the wild, which was negatively related to the light intensity. Our results do not support the growth-mortality trade-off of the POLS hypothesis as highly active, fast-growing individuals showed higher survival than inactive conspecifics. These novel results show for the first time that active and inactive individuals, as scored in the lab, can show different circadian patterns of behavior in the wild driven by light intensity. This implies that studies conducted under a narrow range of light conditions can bias our understanding of individual behavioral variation and its fitness consequences in the wild.

Exploration Behavior and Morphology are Correlated in Captive Gray Mouse Lemurs (Microcebus murinus)

Behavior varies among individuals and is flexible within individuals. However, studies of behavioral syndromes and animal personality have demonstrated that animals can show consistency in their behavior and as such may be restricted in their behavioral responses. Like any other trait, including morphology, performance, or physiology, personality is now considered an important component of ecology and may have fitness consequences. Moreover, in some species personality correlates with other traits, as predicted in the context of a recent theoretical framework postulating that individual differences in growth and body size can affect behavior through effects on growth–mortality tradeoffs. This “pace of life” hypothesis predicts that animals that explore more should be larger and have higher growth rates than those that explore less. We tested for associations between morphology and a behavioral trait in a captive colony of gray mouse lemurs (Microcebus murinus). We used open-field tests to evaluate exploration behavior and measured a series of morphological traits in 72 individuals (32 males and 40 females). Our results show that the latency to start exploring correlates positively with adult body size and body weight at birth. These data provide evidence for a link between morphology and behavior in this species, thus supporting predictions of dispersal models but diverging from the predictions of the “pace of life” model.

A trait-based trade-off between growth and mortality: evidence from 15 tropical tree species using size-specific relative growth rates.

A life-history trade-off between low mortality in the dark and rapid growth in the light is one of the most widely accepted mechanisms underlying plant ecological strategies in tropical forests. Differences in plant functional traits are thought to underlie these distinct ecological strategies; however, very few studies have shown relationships between functional traits and demographic rates within a functional group. We present 8 years of growth and mortality data from saplings of 15 species of Dipterocarpaceae planted into logged-over forest in Malaysian Borneo, and the relationships between these demographic rates and four key functional traits: wood density, specific leaf area (SLA), seed mass, and leaf C:N ratio. Species-specific differences in growth rates were separated from seedling size effects by fitting nonlinear mixed-effects models, to repeated measurements taken on individuals at multiple time points. Mortality data were analyzed using binary logistic regressions in a mixed-effects models framework. Growth increased and mortality decreased with increasing light availability. Species differed in both their growth and mortality rates, yet there was little evidence for a statistical interaction between species and light for either response. There was a positive relationship between growth rate and the predicted probability of mortality regardless of light environment, suggesting that this relationship may be driven by a general trade-off between traits that maximize growth and traits that minimize mortality, rather than through differential species responses to light. Our results indicate that wood density is an important trait that indicates both the ability of species to grow and resistance to mortality, but no other trait was correlated with either growth or mortality. Therefore, the growth mortality trade-off among species of dipterocarp appears to be general in being independent of species crossovers in performance in different light environments.

Individual and sex‐specific differences in intrinsic growth rate covary with consistent individual differences in behaviour

The evolutionary causes of consistent individual differences in behaviour are currently a source of debate. A recent hypothesis suggests that consistent individual differences in life-history productivity (growth and/or fecundity) may covary with behavioural traits that contribute to growth-mortality trade-offs, such as risk-proneness (boldness) and foraging activity (voraciousness). It remains unclear, however, to what extent individual behavioural and life-history profiles are set early in life, or are a more flexible result of specific environmental or developmental contexts that allow bold and active individuals to acquire more resources. Longitudinal studies of individually housed animals under controlled conditions can shed light on this question. Since growth and behaviour can both vary within individuals (they are labile), studying between-individual correlations in behaviour and growth rate requires repeated scoring for both variables over an extended period of time. However, such a study has not yet been done. Here, we repeatedly measured individual mass seven times each, boldness 40 times each and voracity eight times each during the first 4months of life on 90 individually housed crayfish (Cherax destructor). Animals were fed ad libitum, generating a context where individuals can express their intrinsic growth rate (i.e. growth capacity), but in which bold and voracious behaviour is not necessary for high resource acquisition (crayfish can and do hoard food back to their burrow). We show that individuals that were consistently bold over time during the day were also bolder at night, were more voracious and maintained higher growth rates over time than shy individuals. Independent of individual differences, we also observed that males were faster-growing, bolder and more voracious than females. Our findings imply that associations between bold behaviour and fast growth can occur in unlimited food contexts where there is no necessary link between bold behaviour and resource acquisition - offering support for the 'personality-productivity' hypothesis. We suggest future research should study links between consistent individual differences in behaviour and life history under a wider range of contexts, in order to shed light on the role of biotic and abiotic conditions in the strength, direction and stability of their covariance.

Functional traits and the growth–mortality trade‐off in tropical trees

A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals (r2 = 0.69), intermediate for saplings for average growth rates and overall mortality rates (r2 = 0.46), and much weaker for large trees (r2 < or = 0.10). This parallels likely levels of spatial variation in light availability, which is greatest for fast- vs. slow-growing saplings and least for large trees with foliage in the forest canopy. Inherent attributes of species contributing to the trade-off include abilities to disperse, acquire resources, grow rapidly, and tolerate shade and other stresses. There is growing interest in the possibility that functional traits might provide insight into such ecological differences and a growing consensus that seed mass (SM), leaf mass per area (LMA), wood density (WD), and maximum height (H(max)) are key traits among forest trees. Seed mass, LMA, WD, and H(max) are predicted to be small for light-demanding species with rapid growth and mortality and large for shade-tolerant species with slow growth and mortality. Six of these trait-demographic rate predictions were realized for saplings; however, with the exception of WD, the relationships were weak (r2 < 0.1 for three and r2 < 0.2 for five of the six remaining relationships). The four traits together explained 43-44% of interspecific variation in species positions on the growth-mortality trade-off; however, WD alone accounted for > 80% of the explained variation and, after WD was included, LMA and H(max) made insignificant contributions. Virtually the full range of values of SM, LMA, and H(max) occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait-based ecology of tropical forest trees will require consideration of additional traits.

Functional traits and the growth-mortality tradeoff in tropical trees

Functional traits and the growth-mortality tradeoff in tropical trees

Growth-mortality trade-offs along a depth gradient in Cancer borealis

Intertidal and shallow subtidal ecosystems experience steep environmental gradients over short distances. Individual foraging rate, predation risk, and physiologic stress vary along these gradients, resulting in growth-mortality trade-offs with depth. In the summer, Cancer borealis commonly forage in the shallow subtidal in the Gulf of Maine. C. borealis are the favored invertebrate prey of the Herring Gull and the Great Black-backed Gull, which consume 25%–50% of available C. borealis (those in < 1 m water) during each daytime low tide. We investigated three possible explanations for the presence of C. borealis in the risky gull-predation zone. First, we tested whether predation risk in the gull-predation zone was matched at deeper depths by subtidal predators; we found predation risk decreases with depth. Second, we tested whether water temperatures were warmer in the gull-predation zone and whether these warmer temperatures resulted in increased growth rates. We found that, while waters were warmer in the gull-predation zone, crabs grew at similar rates above and below the thermocline when fed similar diets. Finally, we tested for differences in food availability with depth and whether these differences influenced C. borealis growth rates. Our results suggest a growth-mortality trade-off, where increased food availability provides sufficient growth benefit to outweigh the risk of foraging at shallower depths.

Growth‐mortality tradeoffs and ‘personality traits’ in animals

Consistent individual differences in boldness, reactivity, aggressiveness, and other 'personality traits' in animals are stable within individuals but vary across individuals, for reasons which are currently obscure. Here, I suggest that consistent individual differences in growth rates encourage consistent individual differences in behavior patterns that contribute to growth-mortality tradeoffs. This hypothesis predicts that behavior patterns that increase both growth and mortality rates (e.g. foraging under predation risk, aggressive defense of feeding territories) will be positively correlated with one another across individuals, that selection for high growth rates will increase mean levels of potentially risky behavior across populations, and that within populations, faster-growing individuals will take more risks in foraging contexts than slower-growing individuals. Tentative empirical support for these predictions suggests that a growth-mortality perspective may help explain some of the consistent individual differences in behavioral traits that have been reported in fish, amphibians, reptiles, and other animals with indeterminate growth.