Personality and fitness consequences of flight initiation distance and mating behavior in subdominant male reindeer (Rangifer tarandus)

Animal personality has been studied extensively in recent years, yet multidimensionality in tendencies of risk‐related behavior, and the role of such consistency from a mating tactics perspective, is yet to be investigated. We used a semi‐domesticated herd of reindeer (Rangifer tarandus) to examine individual subdominant male propensity to risk mating attempts on guarded females, as well as flight initiation distance (FID), within the personality paradigm to elucidate potential fitness consequences of consistency from an adaptive perspective. Data were collected at the Kutuharju Reindeer Research Station in Kaamanen, Finland, where measures of personality were generated using field observation data based on the relative frequency of dominant male–subdominant male agonistic interactions over 4 years and subdominant males' FID measured over 1 year. Individual propensity for transient mating attempts was not significantly repeatable and did not significantly predict reproductive success or somatic cost during the mating season. Individuals varied consistently in FID, and although repeatable, FID was not related to reproductive success or somatic cost. Proximate state‐dependent or social mechanisms may be driving decision‐making with respect to mating effort, whereas consistent between‐individual differences in FID may be maintained by mechanisms unrelated to life‐history trade‐offs involving productivity.     

Beyond the fast–slow continuum: demographic dimensions structuring a tropical tree community

Life‐history theory posits that trade‐offs between demographic rates constrain the range of viable life‐history strategies. For coexisting tropical tree species, the best established demographic trade‐off is the growth‐survival trade‐off. However, we know surprisingly little about co‐variation of growth and survival with measures of reproduction. We analysed demographic rates from seed to adult of 282 co‐occurring tropical tree and shrub species, including measures of reproduction and accounting for ontogeny. Besides the well‐established fast–slow continuum, we identified a second major dimension of demographic variation: a trade‐off between recruitment and seedling performance vs. growth and survival of larger individuals (≥ 1 cm dbh) corresponding to a ‘stature–recruitment’ axis. The two demographic dimensions were almost perfectly aligned with two independent trait dimensions (shade tolerance and size). Our results complement recent analyses of plant life‐history variation at the global scale and reveal that demographic trade‐offs along multiple axes act to structure local communities.     

Longer life span is associated with elevated immune activity in a seasonally polyphenic butterfly

Seasonal polyphenism constitutes a specific type of phenotypic plasticity in which short‐lived organisms produce different phenotypes in different times of the year. Seasonal generations of such species frequently differ in their overall lifespan and in the values of traits closely related to fitness. Seasonal polyphenisms provide thus excellent, albeit underused model systems for studying trade‐offs between life‐history traits. Here, we compare immunological parameters between the two generations of the European map butterfly (Araschnia levana), a well‐known example of a seasonally polyphenic species. To reveal possible costs of immune defence, we also examine the concurrent differences in several life‐history traits. Both in laboratory experiments and in the field, last instar larvae heading towards the diapause (overwintering) had higher levels of both phenoloxidase (PO) activity and lytic activity than directly developing individuals. These results suggest that individuals from the diapausing generation with much longer juvenile (pupal) period invest more in their immune system than those from the short‐living directly developing generation. The revealed negative correlation between pupal mass and PO activity may be one of the reasons why, in this species, the diapausing generation has a smaller body size than the directly developing generation. Immunological parameters may thus well mediate trade‐offs between body size‐related traits.     

Hematocrit,age, and survival in a wild vertebrate population

Understanding trade‐offs in wild populations is difficult, but important if we are to understand the evolution of life histories and the impact of ecological variables upon them. Markers that reflect physiological state and predict future survival would be of considerable benefit to unraveling such trade‐offs and could provide insight into individual variation in senescence. However, currently used markers often yield inconsistent results. One underutilized measure is hematocrit, the proportion of blood comprising erythrocytes, which relates to the blood's oxygen‐carrying capacity and viscosity, and to individual endurance. Hematocrit has been shown to decline with age in cross‐sectional studies (which may be confounded by selective appearance/disappearance). However, few studies have tested whether hematocrit declines within individuals or whether low hematocrit impacts survival in wild taxa. Using longitudinal data from the Seychelles warbler (Acrocephalus sechellensis), we demonstrated that hematocrit increases with age in young individuals (<1.5 years) but decreases with age in older individuals (1.5–13 years). In breeders, hematocrit was higher in males than females and varied relative to breeding stage. High hematocrit was associated with lower survival in young individuals, but not older individuals. Thus, while we did not find support for hematocrit as a marker of senescence, high hematocrit is indicative of poor condition in younger individuals. Possible explanations are that these individuals were experiencing dehydration and/or high endurance demands prior to capture, which warrants further investigation. Our study demonstrates that hematocrit can be an informative metric for life‐history studies investigating trade‐offs between survival, longevity, and reproduction.     

After the games are over: life‐history trade‐offs drive dispersal attenuation following range expansion

Increased dispersal propensity often evolves on expanding range edges due to the Olympic Village effect, which involves the fastest and fittest finding themselves together in the same place at the same time, mating, and giving rise to like individuals. But what happens after the range's leading edge has passed and the games are over? Although empirical studies indicate that dispersal propensity attenuates following range expansion, hypotheses about the mechanisms driving this attenuation have not been clearly articulated or tested. Here, we used a simple model of the spatiotemporal dynamics of two phenotypes, one fast and the other slow, to propose that dispersal attenuation beyond preexpansion levels is only possible in the presence of trade‐offs between dispersal and life‐history traits. The Olympic Village effect ensures that fast dispersers preempt locations far from the range's previous limits. When trade‐offs are absent, this preemptive spatial advantage has a lasting impact, with highly dispersive individuals attaining equilibrium frequencies that are strictly higher than their introduction frequencies. When trade‐offs are present, dispersal propensity decays rapidly at all locations. Our model's results about the postcolonization trajectory of dispersal evolution are clear and, in principle, should be observable in field studies. We conclude that empirical observations of postcolonization dispersal attenuation offer a novel way to detect the existence of otherwise elusive trade‐offs between dispersal and life‐history traits.     

Effects of variation in resource acquisition during different stages of the life cycle on life‐history traits and trade‐offs in a burying beetle

Individual variation in resource acquisition should have consequences for life‐history traits and trade‐offs between them because such variation determines how many resources can be allocated to different life‐history functions, such as growth, survival and reproduction. Since resource acquisition can vary across an individual's life cycle, the consequences for life‐history traits and trade‐offs may depend on when during the life cycle resources are limited. We tested for differential and/or interactive effects of variation in resource acquisition in the burying beetle Nicrophorus vespilloides. We designed an experiment in which individuals acquired high or low amounts of resources across three stages of the life cycle: larval development, prior to breeding and the onset of breeding in a fully crossed design. Resource acquisition during larval development and prior to breeding affected egg size and offspring survival, respectively. Meanwhile, resource acquisition at the onset of breeding affected size and number of both eggs and offspring. In addition, there were interactive effects between resource acquisition at different stages on egg size and offspring survival. However, only when females acquired few resources at the onset of breeding was there evidence for a trade‐off between offspring size and number. Our results demonstrate that individual variation in resource acquisition during different stages of the life cycle has important consequences for life‐history traits but limited effects on trade‐offs. This suggests that in species that acquire a fixed‐sized resource at the onset of breeding, the size of this resource has larger effects on life‐history trade‐offs than resources acquired at earlier stages.     

Testing the phenotype‐linked fertility hypothesis in the presence and absence of inbreeding

The phenotype‐linked fertility hypothesis suggests that females can judge male fertility by inspecting male phenotypic traits. This is because male sexually selected traits might correlate with sperm quality if both are sensitive to factors that influence male condition. A recent meta‐analysis found little support for this hypothesis, suggesting little or no shared condition dependence. However, we recently reported that in captive zebra finches (Taeniopygia guttata) inbreeding had detrimental effects both on phenotypic traits and on measures of sperm quality, implying that variation in inbreeding could induce positive covariance between indicator traits and sperm quality. Therefore, we here assess empirically the average strength of correlations between phenotypic traits (courtship rate, beak colour, tarsus length) and measures of sperm quality (proportion of functional sperm, sperm velocity, sperm length) in populations of only outbred individuals and in mixed populations consisting of inbreds (F = 0.25) and outbreds (F = 0). As expected, phenotype sperm‐trait correlations were stronger when the population contained a mix of inbred and outbred individuals. We also found unexpected heterogeneity between our two study populations, with correlations being considerably stronger in a domesticated population than in a recently wild‐derived population. Correlations ranged from essentially zero among outbred‐only wild‐derived birds (mean Fisher's Zr ± SE = 0.03 ± 0.10) to moderately strong among domesticated birds of mixed inbreeding status (Zr ± SE = 0.38 ± 0.08). Our results suggest that, under some conditions, the phenotype‐linked fertility hypothesis might apply.     

Life‐history trade‐offs vary with resource availability across the geographic range of a widespread plant

• Trade‐offs between reproduction, growth and survival arise from limited resource availability in plants. Environmental stress is expected to exacerbate these negative correlations, but no studies have evaluated variation in life‐history trade‐offs throughout species geographic ranges. Here we analyse the costs of growth and reproduction across the latitudinal range of the widespread herb Plantago coronopus in Europe.
• We monitored the performance of thousands of individuals in 11 populations of P. coronopus, and tested whether the effects of growth and reproduction on a set of vital rates (growth, probability of survival, probability of reproduction and fecundity) varied with local precipitation and soil fertility. To account for variation in internal resources among individuals, we analysed trade‐offs correcting for differences in size.
• Growth was negatively affected by previous growth and reproduction. We also found costs of growth and reproduction on survival, reproduction probability and fecundity, but only in populations with low soil fertility. Costs also increased with precipitation, possibly due to flooding‐related stress. In contrast, growth was positively correlated with subsequent survival, and there was a positive covariation in reproduction between consecutive years under certain environments, a potential strategy to exploit temporary benign conditions.
• Overall, we found both negative and positive correlations among vital rates across P. coronopus geographic range. Trade‐offs predominated under stressful conditions, and positive correlations arose particularly between related traits like reproduction investment across years. By analysing multiple and diverse fitness components along stress gradients, we can better understand life‐history evolution across species’ ranges, and their responses to environmental change.

     

Flight initiation distance in Lepidopterans is species-specific and positively related to starting distance

Escape behaviour is a critical component of invertebrate life history but is poorly studied. Flight initiation distance (FID) indexes escape propensity, and is well-studied in vertebrates but is entirely unstudied in Lepidopterans, despite their obvious escape behaviour. Here we test two general principles regarding FID as derived from studies of vertebrates to examine if they apply to Sri Lankan butterflies: 1) that FID is a species-specific trait and 2) that FID increases with Starting Distance, the distance at which the experimenter begins an approach. We collected 295 FIDs from 17 species and find that 1) FIDs are a tractable way of indexing butterfly escape and 2) both the general principles tested apply to butterfly escape. We also present FIDs of these species to encourage further data collection and comparative analysis of butterfly escape.     

STRONG SELECTION GENOME‐WIDE ENHANCES FITNESS TRADE‐OFFS ACROSS ENVIRONMENTS AND EPISODES OF SELECTION

Fitness trade‐offs across episodes of selection and environments influence life‐history evolution and adaptive population divergence. Documenting these trade‐offs remains challenging as selection can vary in magnitude and direction through time and space. Here, we evaluate fitness trade‐offs at the levels of the whole organism and the quantitative trait locus (QTL) in a multiyear field study of Boechera stricta (Brassicaceae), a genetically tractable mustard native to the Rocky Mountains. Reciprocal local adaptation was pronounced for viability, but not for reproductive components of fitness. Instead, local genomes had a fecundity advantage only in the high latitude garden. By estimating realized selection coefficients from individual‐level data on viability and reproductive success and permuting the data to infer significance, we examined the genetic basis of fitness trade‐offs. This analytical approach (Conditional Neutrality‐Antagonistic Pleiotropy, CNAP) identified genetic trade‐offs at a flowering phenology QTL (costs of adaptation) and revealed genetic trade‐offs across fitness components (costs of reproduction). These patterns would not have emerged from traditional ANOVA‐based QTL mapping. Our analytical framework can be applied to other systems to investigate fitness trade‐offs. This task is becoming increasingly important as climate change may alter fitness landscapes, potentially disrupting fitness trade‐offs that took many generations to evolve.     

Early‐life telomere length predicts lifespan and lifetime reproductive success in a wild bird

Poor conditions during early development can initiate trade‐offs that favour current survival at the expense of somatic maintenance and subsequently, future reproduction. However, the mechanisms that link early and late life‐history are largely unknown. Recently it has been suggested that telomeres, the nucleoprotein structures at the terminal end of chromosomes, could link early‐life conditions to lifespan and fitness. In wild purple‐crowned fairy‐wrens, we combined measurements of nestling telomere length (TL) with detailed life‐history data to investigate whether early‐life TL predicts fitness prospects. Our study differs from previous studies in the completeness of our fitness estimates in a highly philopatric population. The association between TL and survival was age‐dependent with early‐life TL having a positive effect on lifespan only among individuals that survived their first year. Early‐life TL was not associated with the probability or age of gaining a breeding position. Interestingly, early‐life TL was positively related to breeding duration, contribution to population growth and lifetime reproductive success because of their association with lifespan. Thus, early‐life TL, which reflects growth, accumulated early‐life stress and inherited TL, predicted fitness in birds that reached adulthood but not noticeably among fledglings. These findings suggest that a lack of investment in somatic maintenance during development particularly affects late life performance. This study demonstrates that factors in early‐life are related to fitness prospects through lifespan, and suggests that the study of telomeres may provide insight into the underlying physiological mechanisms linking early‐ and late‐life performance and trade‐offs across a lifetime.     

Social immunity in honeybees—Density dependence,diet, and body mass trade‐offs

Group living is favorable to pathogen spread due to the increased risk of disease transmission among individuals. Similar to individual immune defenses, social immunity, that is antiparasite defenses mounted for the benefit of individuals other than the actor, is predicted to be altered in social groups. The eusocial honey bee (Apis mellifera) secretes glucose oxidase (GOX), an antiseptic enzyme, throughout its colony, thereby providing immune protection to other individuals in the hive. We conducted a laboratory experiment to investigate the effects of group density on social immunity, specifically GOX activity, body mass and feeding behavior in caged honey bees. Individual honeybees caged in a low group density displayed increased GOX activity relative to those kept at a high group density. In addition, we provided evidence for a trade‐off between GOX activity and body mass: Individuals caged in the low group density had a lower body mass, despite consuming more food overall. Our results provide the first experimental evidence that group density affects a social immune response in a eusocial insect. Moreover, we showed that the previously reported trade‐off between immunity and body mass extends to social immunity. GOX production appears to be costly for individuals, and potentially the colony, given that low body mass is correlated with small foraging ranges in bees. At high group densities, individuals can invest less in social immunity than at low densities, while presumably gaining shared protection from infection. Thus, there is evidence that trade‐offs at the individual level (GOX vs. body mass) can affect colony‐level fitness.     

The cost of defense in social insects: insights from the honey bee

Defense is one of the most important factors affecting life history. The relationship of defense to life history traits as well as its possible costs has been reviewed extensively for many groups, including plants. However, defense in social insects, such as honey bees, has never been examined from a trade‐off perspective, although defense in honey bees, Apis mellifera L. (Hymenoptera: Apidae), has been widely studied. In this review, we discuss the life history traits of honey bees, particularly traits related to defense. We then examine trade‐offs in the context of resource availability. Lastly, we offer suggestions for future research on trade‐offs in honey bees and other social insects.     

The development of a synthetic diet for investigating the effects of macronutrients on the development of Plodia interpunctella

The use of chemically defined artificial diets has allowed researchers to examine questions within nutritional ecology about how macronutrients affect life‐history traits and resource‐based trade‐offs. Using a chemically defined diet, it is possible to manipulate both the total nutritional content and the ratio of macronutrients (i.e., proteins, carbohydrates, or lipids) within the diet. Studies using the geometric framework have made use of these diets to examine lifespan, fecundity, and immune responses. Here, we develop an artificial diet suitable for rearing lepidopteran larvae. We created diets with three proportions of non‐nutritive material (30, 50, and 70% indigestible cellulose) relative to protein and carbohydrate macronutrients, and compared these to standard wheat bran laboratory diet. We then examined the effects of variable nutrient content on lifespan and development time in Plodia interpunctella Hübner (Lepidoptera: Pyralidae). The artificial diets supported development (almost) as well as bran‐based laboratory diets. Total nutrient content affected development time: females that fed on the diet with the highest nutrient content took the longest time to reach eclosion. We also found evidence to support dietary restriction, with larvae receiving the fewest nutrients having the longest lifespan as adults. These findings are indicative of the usefulness of this diet as a tool to further investigate the effects of nutrient content and macronutrient imbalance on resource‐based trade‐offs and life‐history traits.     

Equal nonbreeding period survival in adults and juveniles of a long‐distant migrant bird

In migrant birds, survival estimates for the different life‐history stages between fledging and first breeding are scarce. First‐year survival is shown to be strongly reduced compared with annual survival of adult birds. However, it remains unclear whether the main bottleneck in juvenile long‐distant migrants occurs in the postfledging period within the breeding ranges or en route. Quantifying survival rates during different life‐history stages and during different periods of the migration cycle is crucial to understand forces driving the evolution of optimal life histories in migrant birds. Here, we estimate survival rates of adult and juvenile barn swallows (Hirundo rustica L.) in the breeding and nonbreeding areas using a population model integrating survival estimates in the breeding ranges based on a large radio‐telemetry data set and published estimates of demographic parameters from large‐scale population‐monitoring projects across Switzerland. Input parameters included the country‐wide population trend, annual productivity estimates of the double‐brooded species, and year‐to‐year survival corrected for breeding dispersal. Juvenile survival in the 3‐week postfledging period was low (S = 0.32; SE = 0.05), whereas in the rest of the annual cycle survival estimates of adults and juveniles were similarly high (S > 0.957). Thus, the postfledging period was the main survival bottleneck, revealing the striking result that nonbreeding period mortality (including migration) is not higher for juveniles than for adult birds. Therefore, focusing future research on sources of variation in postfledging mortality can provide new insights into determinants of population dynamics and life‐history evolution of migrant birds.     

The trade‐off between fecundity and egg size in a polymorphic population of Arctic charr (Salvelinus alpinus (L.)) in Skogsfjordvatn,subarctic Norway

Reproductive traits differ between intralacustrine Arctic charr morphs. Here, we examine three sympatric lacustrine Arctic charr morphs with respect to fecundity, egg size and spawning time/site to assess reproductive investments and trade‐offs, and possible fitness consequences. The littoral omnivore morph (LO‐morph) utilizes the upper water for feeding and reproduction and spawn early in October. The large profundal piscivore morph (PP‐morph) and the small profundal benthivore morph (PB‐morph) utilize the profundal habitat for feeding and reproduction and spawn in December and November, respectively. Females from all morphs were sampled for fecundity and egg‐size analysis. There were large differences between the morphs. The PB‐morph had the lowest fecundity (mean = 45, SD = 13) and smallest egg size (mean = 3.2 mm, SD = 0.32 mm). In contrast, the PP‐morph had the highest fecundity (mean = 859.5, SD = 462) and the largest egg size (mean = 4.5 mm, SD = 0.46 mm), whereas the LO‐morph had intermediate fecundity (mean = 580, SD = 225) and egg size (mean = 4.3, SD = 0.24 mm). Fecundity increased with increasing body size within each morph. This was not the case for egg size, which was independent of body sizes within morph. Different adaptations to feeding and habitat utilization have apparently led to a difference in the trade‐off between fecundity and egg size among the three different morphs.     

THE EFFECTS OF PREDATION RISK ON MATING SYSTEM EXPRESSION IN A FRESHWATER SNAIL

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

Hot and not‐so‐hot females: reproductive state and thermal preferences of female Arizona Bark Scorpions (Centruroides sculpturatus)

For ectotherms, environmental temperatures influence numerous life history characteristics, and the body temperatures (T_b) selected by individuals can affect offspring fitness and parental survival. Reproductive trade‐offs may therefore ensue for gravid females, because temperatures conducive to embryonic development may compromise females' body condition. We tested whether reproduction influenced thermoregulation in female Arizona Bark Scorpions (Centruroides sculpturatus). We predicted that gravid females select higher T_b and thermoregulate more precisely than nonreproductive females. Gravid C. sculpturatus gain body mass throughout gestation, which exposes larger portions of their pleural membrane, possibly increasing their rates of transcuticular water loss in arid environments. Accordingly, we tested whether gravid C. sculpturatus lose water faster than nonreproductive females. We determined the preferred T_b of female scorpions in a thermal gradient and measured water loss rates using flow‐through respirometry. Gravid females preferred significantly higher T_b than nonreproductive females, suggesting that gravid C. sculpturatus alter their thermoregulatory behaviour to promote offspring fitness. However, all scorpions thermoregulated with equal precision, perhaps because arid conditions create selective pressure on all females to thermoregulate effectively. Gravid females lost water faster than nonreproductive animals, indicating that greater exposure of the pleural membrane during gestation enhances the desiccation risk of reproductive females. Our findings suggest that gravid C. sculpturatus experience a trade‐off, whereby selection of higher T_b and increased mass during gestation increase females' susceptibility to water loss, and thus their mortality risk. Elucidating the mechanisms that influence thermal preferences may reveal how reproductive trade‐offs shape the life history of ectotherms in arid environments.     

Effects of pathogen exposure on life‐history variation in the gypsy moth (Lymantria dispar)

Investment in host defences against pathogens may lead to trade‐offs with host fecundity. When such trade‐offs arise from genetic correlations, rates of phenotypic change by natural selection may be affected. However, genetic correlations between host survival and fecundity are rarely quantified. To understand trade‐offs between immune responses to baculovirus exposure and fecundity in the gypsy moth (Lymantria dispar), we estimated genetic correlations between survival probability and traits related to fecundity, such as pupal weight. In addition, we tested whether different virus isolates have different effects on male and female pupal weight. To estimate genetic correlations, we exposed individuals of known relatedness to a single baculovirus isolate. To then evaluate the effect of virus isolate on pupal weight, we exposed a single gypsy moth strain to 16 baculovirus isolates. We found a negative genetic correlation between survival and pupal weight. In addition, virus exposure caused late‐pupating females to be identical in weight to males, whereas unexposed females were 2–3 times as large as unexposed males. Finally, we found that female pupal weight is a quadratic function of host mortality across virus isolates, which is likely due to trade‐offs and compensatory growth processes acting at high and low mortality levels, respectively. Overall, our results suggest that fecundity costs may strongly affect the response to selection for disease resistance. In nature, baculoviruses contribute to the regulation of gypsy moth outbreaks, as pathogens often do in forest‐defoliating insects. We therefore argue that trade‐offs between host life‐history traits may help explain outbreak dynamics.     

When do trade‐offs occur? The roles of energy constraints and trait flexibility in bushcricket populations

In many animal species, the expression of sexually selected traits is negatively correlated with traits associated with survival such as immune function, a relationship termed a ‘trade‐off’. But an alternative in which sexually selected traits are positively correlated with survival traits is also widespread. We propose that the nature of intertrait relationships is largely determined by overall energy expenditure, energy availability and trait flexibility, with trade‐offs expected when individuals are subject to energy constraints. We tested this hypothesis in Ephippiger diurnus, a European bushcricket in which males are distinguished by two prominent sexually selected traits, acoustic calls and a large spermatophore transferred to the female at mating, and where immune function may be critical in survival. Ephippiger diurnus are distributed as small, isolated populations that are differentiated genetically and behaviourally. We analysed songs, spermatophores and the immune function in male individuals from eight populations spanning a range of song types. As predicted, we only found trade‐offs in those populations that expended more energy on song and were less flexible in their ability to adjust that expenditure. Ultimately, energy constraints and resulting trade‐offs may limit the evolution of song exaggeration in E. diurnus populations broadcasting long calls comprised of multiple ‘syllables’.