RESUMO
Early-life adversity, even when transient, can have lasting effects on individual phenotypes and reduce lifespan across species. If these effects can be mitigated by a high-quality later-life environment, then differences in future resources may explain variable resilience to early-life adversity. Using data from over 1000 wild North American red squirrels, we tested the hypothesis that the costs of early-life adversity for adult lifespan could be offset by later-life food abundance. We identified six adversities that reduced juvenile survival in the first year of life, though only one-birth date-had continued independent effects on adult lifespan. We then built a weighted early-life adversity (wELA) index integrating the sum of adversities and their effect sizes. Greater weighted early-life adversity predicted shorter adult lifespans in males and females, but a naturally occurring food boom in the second year of life ameliorated this effect. Experimental food supplementation did not replicate this pattern, despite increasing lifespan, indicating that the buffering effect of a future food boom may hinge on more than an increase in available calories. Our results suggest a non-deterministic role of early-life conditions for later-life phenotype, highlighting the importance of evaluating the consequences of early-life adversity in the context of an animal's entire life course.
Assuntos
Longevidade , Sciuridae , Animais , Masculino , Feminino , Sciuridae/fisiologiaRESUMO
Many studies assume that it is beneficial for individuals of a species to be heavier, or have a higher body condition index (BCI), without accounting for the physiological relevance of variation in the composition of different body tissues. We hypothesized that the relationship between BCI and masses of physiologically important tissues (fat and lean) would be conditional on annual patterns of energy acquisition and expenditure. We studied three species with contrasting ecologies in their respective natural ranges: an obligate hibernator (Columbian ground squirrel, Urocitellus columbianus), a facultative hibernator (black-tailed prairie dog, Cynomys ludovicianus), and a food-caching non-hibernator (North American red squirrel, Tamiasciurus hudsonicus). We measured fat and lean mass in adults of both sexes using quantitative magnetic resonance (QMR). We measured body mass and two measures of skeletal structure (zygomatic width and right hind foot length) to develop sex- and species-specific BCIs, and tested the utility of BCI to predict body composition in each species. Body condition indices were more consistently, and more strongly correlated, with lean mass than fat mass. The indices were most positively correlated with fat when fat was expected to be very high (pre-hibernation prairie dogs). In all cases, however, BCI was never better than body mass alone in predicting fat or lean mass. While the accuracy of BCI in estimating fat varied across the natural histories and annual energetic patterns of the species considered, measuring body mass alone was as effective, or superior in capturing sufficient variation in fat and lean in most cases.
Assuntos
Composição Corporal , Alimentos , Humanos , Masculino , Feminino , Animais , Composição Corporal/fisiologia , Sciuridae/fisiologia , Especificidade da EspécieRESUMO
While cooperative interactions among kin are a key building block in the societies of group-living species, their importance for species with more variable social environments is unclear. North American red squirrels (Tamiasciurus hudsonicus) defend individual territories in dynamic neighbourhoods and are known to benefit from living among familiar conspecifics, but not relatives. However, kin-directed behaviours may be restricted to specific genealogical relationships or strongly mediated by geographical distance, masking their influence at broader scales. Using distance between territories as a proxy for the ability of individuals to interact, we estimated the influence of primary kin (parents, offspring, siblings) on the annual survival and reproductive success of red squirrels. This approach revealed associations between fitness and access to kin, but only for certain genealogical relationships and fitness components. For example, females had enhanced annual survival when living closer to their daughters, though the reverse was not true. Most surprising was the finding that males had higher annual reproductive success when living closer to their father, suggesting possible recognition and cooperation among fathers and sons. Together, these findings point to unexpected nuance in the fitness consequences of kinship dynamics for a species that is territorial and largely solitary.
Assuntos
Irmãos , Territorialidade , Humanos , Animais , Masculino , Feminino , Sciuridae , Reprodução , Meio Social , Comportamento SocialRESUMO
Desynchrony of phenological responses to climate change is a major concern for ecological communities. Potential uncoupling between one of the most fundamental divisions within populations, males and females, has not been well studied. To address this gap, we examined sex-specific plasticity in hibernation phenology in two populations of Columbian ground squirrels (Urocitellus columbianus). We find that both sexes display similar phenological plasticity to spring snowmelt dates in their timing of torpor termination and behavioural emergence from hibernation. As a result of this plasticity, the degree of protandry (i.e. males' emergences from hibernation preceding those of females) did not change significantly over the 27-year study. Earlier male behavioural emergence, relative to females, improved the likelihood of securing a breeding territory and increased annual reproductive success. Sexual selection favouring earlier male emergence from hibernation may maintain protandry in this population, but did not contribute to further advances in male phenology. Together, our results provide evidence that the sexes should remain synchronized, at least in response to the weather variation investigated here, and further support the role of sexual selection in the evolution of protandry in sexually reproducing organisms.
Assuntos
Sexo , Seleção Sexual , Feminino , Animais , Masculino , Reprodução/fisiologia , Adaptação Fisiológica , Estações do Ano , Sciuridae/fisiologiaRESUMO
Territories are typically defined as spatially exclusive areas that are defended against conspecifics. Given the spatial nature of territoriality, it is inherently density dependent, but the economics of territoriality also depend on the distribution and abundance of defended resources. Our objectives were to assess the effects of changing population density and food availability on individually based territorial phenotypes. We developed a novel analytical framework that bridges spatially explicit territories with social network analysis to model density-dependent territorial phenotypes. Using the outputs from our data pipeline, we modelled plasticity in territory size and territory intrusion rates in a long-term study population of North American red squirrels Tamiasciurus hudsonicus. Red squirrels defend year-round territories around a central hoard (midden) of white spruce Picea glauca cones. Importantly, white spruce is a masting species that produces large cone crops every 4-7 years (i.e. mast years) in our study area interspersed with non-mast years when few cones are produced. In the spring following mast years, populations are approximately double in size, but are lower in the spring of non-mast years. We predicted that territory size and intrusion rates would decrease as resource abundance, and consequently population density, increased. By contrast, as resource abundance decreased via depletion, and therefore density decreased, territories should increase in size and intrusions should also increase. As we expected, individual territory size and territorial intrusions were negatively density dependent, such that increased density after mast years resulted in smaller territories and fewer intrusions. When considering between-individual variation in plasticity across a density gradient, individuals responded differently to changes in population density within their lifetime. Our results show that territory size and intrusion rates display negative density dependence. When food becomes available in the autumn of a mast year and density in spring of the following year increases, territories shrink in size to effectively a small area around the midden. While our findings for red squirrels are unique compared to other systems, they serve as a reminder that the direction and strength of fundamental ecological relationships can depend on the nature of the system.
Assuntos
Análise de Rede Social , Territorialidade , Animais , Densidade Demográfica , SciuridaeRESUMO
INTRODUCTION: Backcountry skiers and snowboarders are increasingly using avalanche airbags to improve safety. New safety devices can cause risk compensation, the concept in which users take more risks given the larger safety margin provided by the device. This may limit overall benefits. We sought to elucidate attitudes toward risk-taking behaviors and risk compensation in backcountry users relating to avalanche airbags. METHODS: A convenience sample of 144 backcountry skiers and snowboarders was surveyed after a backcountry tour in the Wasatch Mountains of Utah and the Tetons of Wyoming during the winter 2020-21 season. Demographic and experiential data were compared with risk propensity scores and attitudes toward risk compensation. Respondents were stratified into high-, medium-, and low-risk groupings based on risk propensity scores and whether an airbag was carried. RESULTS: Thirty-two (22%) respondents carried an airbag. Airbag users were more likely to endorse risk compensation behavior, ski terrain over 30 degrees, and fall into the high-risk cohort. The high-risk cohort was also more likely to endorse risk compensation behavior than medium- and low-risk individuals. CONCLUSIONS: Risk compensation was more prevalent in 2 groups: 1) those carrying an airbag and 2) those falling within the high-risk cohort. Given the prevalence of avalanche airbags, risk compensation should be considered alongside other human factors in avalanche safety and education so that users can mitigate these effects. Although risk compensation appears to be occurring, the magnitude of this effect remains unknown and likely does not obviate the safety benefits of the airbag altogether.
Assuntos
Avalanche , Esqui , Humanos , Esqui/educação , Equipamentos de Proteção , Utah , AtitudeRESUMO
The Cort-Adaptation hypothesis suggests that elevated glucocorticoids (GCs) can facilitate an adaptive response to environmental and physiological challenges. Most previous studies have focused on avian species, which may limit their generalizability to mammals, where lactation is known to be a major physiological challenge. Furthermore, the effect of predation risk on GC levels has not been tested in the Cort-Adaptation hypothesis. We sought to test this hypothesis in a colonial prey species, black-tailed prairie dogs (Cynomys ludovicianus). We predicted that individuals located near fewer neighboring conspecifics would perceive an increased risk of predation and, in turn, have increased GCs (measured through hair cortisol concentration (HCC)) and reduced annual reproductive success compared to more centrally located individuals. We also investigated other putative influences on HCC: age, lactation status, body condition, and season of hair growth. Levels of vigilance behavior were higher for those with fewer neighboring conspecifics, suggesting variation in perceived risk of predation. Further, the risk of predation appeared to represent a chronic, detrimental stressor as evidenced by a significant increase in HCC for prairie dogs with fewer neighbors. Lactation status and season also influenced HCC. We found support for the Cort-Adaptation hypothesis where increased HCC during the reproductive season correlated with whether a female produced a litter, but not litter size, suggesting a minimum threshold of GCs is required for successful reproduction in this species. Our work illustrates that HCC may operate as an indicator of perceived predation risk, but care should be taken to consider the variety of factors influencing GC homeostasis, in particular lactation, when drawing conclusions using HCC as a marker of long-term stress.
Assuntos
Glucocorticoides , Comportamento Predatório , Animais , Feminino , Hidrocortisona , Reprodução/fisiologia , SciuridaeRESUMO
Evolutionary endocrinology aims to understand how natural selection shapes endocrine systems and the degree to which endocrine systems themselves can induce phenotypic responses to environmental changes. Such responses may be specialized in that they reflect past selection for responsiveness only to those ecological factors that ultimately influence natural selection. Alternatively, endocrine responses may be broad and generalized, allowing organisms to cope with a variety of environmental changes simultaneously. Here, we empirically tested whether the endocrine response of female North American red squirrels (Tamiasciurus hudsonicus) was specialized or generalized. We first quantified the direction and magnitude of natural selection acting on three female life history traits (parturition date, litter size, offspring postnatal growth rate) during 32 years of fluctuations in four potential ecological agents of selection (food availability, conspecific density, predator abundance, and temperature). Only three of the four variables (food, density, and predators) affected patterns of natural selection on female life history traits. We then quantified fecal glucocorticoid metabolites (FGMs) across 7 years and found that all four environmental variables, regardless of their effects on patterns of selection, were associated with glucocorticoid production. Our results provide support for a generalized, rather than specific, glucocorticoid response to environmental change that can integrate across multiple co-occurring environmental stressors.
Assuntos
Glucocorticoides , Seleção Genética , Animais , Gravidez , Feminino , Sciuridae/fisiologia , Evolução Biológica , Tamanho da Ninhada de Vivíparos/fisiologiaRESUMO
When resources are limited, mean fitness is constrained and competition can cause genes and phenotypes to enhance an individual's own fitness while reducing the fitness of their competitors. Negative social effects on fitness have the potential to constrain adaptation, but the interplay between ecological opportunity and social constraints on adaptation remains poorly studied in nature. Here, we tested for evidence of phenotypic social effects on annual fitness (survival and reproductive success) in a long-term study of wild North American red squirrels (Tamiasciurus hudsonicus) under conditions of both resource limitation and super-abundant food resources. When resources were limited, populations remained stable or declined, and there were strong negative social effects on annual survival and reproductive success. That is, mean fitness was constrained and individuals had lower fitness when other nearby individuals had higher fitness. In contrast, when food resources were super-abundant, populations grew and social constraints on reproductive success were greatly reduced or eliminated. Unlike reproductive success, social constraints on survival were not significantly reduced when food resources were super-abundant. These findings suggest resource-dependent social constraints on a component of fitness, which have important potential implications for evolution and adaptation.
Assuntos
Reprodução , Sciuridae , Adaptação Fisiológica , Animais , Fenótipo , Sciuridae/genéticaRESUMO
As a response to environmental cues, maternal glucocorticoids (GCs) may trigger adaptive developmental plasticity in the physiology and behavior of offspring. In North American red squirrels (Tamiasciurus hudsonicus), mothers exhibit increased GCs when conspecific density is elevated, and selection favors more aggressive and perhaps more active mothers under these conditions. We tested the hypothesis that elevated maternal GCs cause shifts in offspring behavior that may prepare them for high-density conditions. We experimentally elevated maternal GCs during gestation or early lactation. We measured two behavioral traits (activity and aggression) in weaned offspring using standardized behavioral assays. Because maternal GCs may influence offspring hypothalamic-pituitary-adrenal (HPA) axis dynamics, which may in turn affect behavior, we also measured the impact of our treatments on offspring HPA axis dynamics (adrenal reactivity and negative feedback), and the association between offspring HPA axis dynamics and behavior. Increased maternal GCs during lactation, but not gestation, slightly elevated activity levels in offspring. Offspring aggression and adrenal reactivity did not differ between treatment groups. Male, but not female, offspring from mothers treated with GCs during pregnancy exhibited stronger negative feedback compared with those from control mothers, but there were no differences in negative feedback between lactation treatment groups. Offspring with higher adrenal reactivity from mothers treated during pregnancy (both controls and GC-treated) exhibited lower aggression and activity. These results suggest that maternal GCs during gestation or early lactation alone may not be a sufficient cue to produce substantial changes in behavioral and physiological stress responses in offspring in natural populations.
Assuntos
Sistema Hipotálamo-Hipofisário , Sistema Hipófise-Suprarrenal , Animais , Feminino , Glucocorticoides , Humanos , Masculino , Comportamento Materno , Gravidez , Sciuridae , Estresse Fisiológico , Estados UnidosRESUMO
Dispersal is nearly universal; yet, which sex tends to disperse more and their success thereafter depends on the fitness consequences of dispersal. We asked if lifetime fitness differed between residents and immigrants (successful between-population dispersers) and their offspring using 29 years of monitoring from North American red squirrels (Tamiasciurus hudsonicus) in Canada. Compared to residents, immigrant females had 23% lower lifetime breeding success (LBS), while immigrant males had 29% higher LBS. Male immigration and female residency were favoured. Offspring born to immigrants had 15-43% lower LBS than offspring born to residents. We conclude that immigration benefitted males, but not females, which appeared to be making the best of a bad lot. Our results are in line with male-biased dispersal being driven by local mate competition and local resource enhancement, while the intergenerational cost to immigration is a new complication in explaining the drivers of sex-biased dispersal.
Assuntos
Emigrantes e Imigrantes , Reprodução , Cruzamento , Canadá , Feminino , Humanos , MasculinoRESUMO
Elevations in glucocorticoid (GC) levels in breeding females may induce adaptive shifts in offspring life histories. Offspring produced by mothers with elevated GCs may be better prepared to face harsh environments, where a faster pace of life is beneficial. We examined how experimentally elevated GCs in pregnant or lactating North American red squirrels (Tamiasciurus hudsonicus) affected offspring postnatal growth, structural size and oxidative stress levels (two antioxidants and oxidative protein damage) in three different tissues (blood, heart and liver) and liver telomere lengths. We predicted that offspring from mothers treated with GCs would grow faster but would also have higher levels of oxidative stress and shorter telomeres, which may predict reduced longevity. Offspring from mothers treated with GCs during pregnancy were 8.3% lighter around birth but grew (in body mass) 17.0% faster than those from controls, whereas offspring from mothers treated with GCs during lactation grew 34.8% slower than those from controls and did not differ in body mass around birth. Treating mothers with GCs during pregnancy or lactation did not alter the oxidative stress levels or telomere lengths of their offspring. Fast-growing offspring from any of the treatment groups did not have higher oxidative stress levels or shorter telomere lengths, indicating that offspring that grew faster early in life did not exhibit oxidative costs after this period of growth. Our results indicate that elevations in maternal GCs may induce plasticity in offspring growth without long-term oxidative costs to the offspring that might result in a shortened lifespan.
Assuntos
Glucocorticoides/metabolismo , Estresse Oxidativo , Sciuridae/fisiologia , Encurtamento do Telômero , Animais , Feminino , Masculino , Sciuridae/crescimento & desenvolvimentoRESUMO
Long-term studies of wild animals provide the opportunity to investigate how phenotypic plasticity is used to cope with environmental fluctuations and how the relationships between phenotypes and fitness can be dependent upon the ecological context. Many previous studies have only investigated life-history plasticity in response to changes in temperature, yet wild animals often experience multiple environmental fluctuations simultaneously. This requires field experiments to decouple which ecological factor induces plasticity in fitness-relevant traits to better understand their population-level responses to those environmental fluctuations. For the past 32 years, we have conducted a long-term integrative study of individually marked North American red squirrels Tamiasciurus hudsonicus Erxleben in the Yukon, Canada. We have used multi-year field experiments to examine the physiological and life-history responses of individual red squirrels to fluctuations in food abundance and conspecific density. Our long-term observational study and field experiments show that squirrels can anticipate increases in food availability and density, thereby decoupling the usual pattern where animals respond to, rather than anticipate, an ecological change. As in many other study systems, ecological factors that can induce plasticity (such as food and density) covary. However, our field experiments that manipulate food availability and social cues of density (frequency of territorial vocalizations) indicate that increases in social (acoustic) cues of density in the absence of additional food can induce similar life-history plasticity, as does experimental food supplementation. Changes in the levels of metabolic hormones (glucocorticoids) in response to variation in food and density are one mechanism that seems to induce this adaptive life-history plasticity. Although we have not yet investigated the energetic response of squirrels to elevated density or its association with life-history plasticity, energetics research in red squirrels has overturned several standard pillars of knowledge in physiological ecology. We show how a tractable model species combined with integrative studies can reveal how animals cope with resource fluctuations through life-history plasticity.
Assuntos
Animais Selvagens , Cauda , Animais , Canadá , Sciuridae , Estados Unidos , YukonRESUMO
Juvenile survival to first breeding is a key life-history stage for all taxa. Survival through this period can be particularly challenging when it coincides with harsh environmental conditions such as a winter climate or food scarcity, leading to highly variable cohort survival. However, the small size and dispersive nature of juveniles generally make studying their survival more difficult. In territorial species, a key life-history event is the acquisition of a territory. A territory is expected to enhance survival, but how it does so is not often identified. We tested how the timing of territory acquisition influenced the winter survival of juvenile North American red squirrels Tamiasciurus hudsonicus, hereafter red squirrels, and how the timing of this event mediated the sources of mortality. We hypothesized that securing a territory prior to when food resources become available would reduce juvenile susceptibility to predation and climatic factors overwinter. Using 27 years of data on the survival of individually marked juvenile red squirrels, we tested how the timing of territory acquisition influenced survival, whether the population density of red squirrel predators and mean temperature overwinter were related to individual survival probability, and if territory ownership mediated these effects. Juvenile red squirrel survival was lower in the years of high predator abundance and in colder winters. Autumn territory owners were less susceptible to lynx Lynx canadensis and possibly mustelid Mustela and Martes spp., predation. Autumn territory owners had lower survival in colder winters, but surprisingly non-owners had higher survival in cold winters. Our results show how the timing of a life-history event like territory acquisition can directly affect survival and also mediate the effects of biotic and abiotic factors later in life. This engenders a better understanding of the fitness consequences of the timing of key life-history events.
Assuntos
Lynx , Sciuridae , Animais , Densidade Demográfica , Comportamento Predatório , Estações do AnoRESUMO
Interactions between organisms are ubiquitous and have important consequences for phenotypes and fitness. Individuals can even influence those they never meet, if they have extended phenotypes that alter the environments others experience. North American red squirrels (Tamiasciurus hudsonicus) guard food hoards, an extended phenotype that typically outlives the individual and is usually subsequently acquired by non-relatives. Hoarding by previous owners can, therefore, influence subsequent owners. We found that red squirrels breed earlier and had higher lifetime fitness if the previous hoard owner was a male. This was driven by hoarding behaviour, as males and mid-aged squirrels had the largest hoards, and these effects persisted across owners, such that if the previous owner was male or died in mid-age, subsequent occupants had larger hoards. Individuals can, therefore, influence each other's resource-dependent traits and fitness without ever meeting, such that the past can influence contemporary population dynamics through extended phenotypes.
Assuntos
Aptidão Genética , Sciuridae , Animais , Masculino , FenótipoRESUMO
Phenological shifts are the most commonly reported ecological responses to climate change and can be produced rapidly by phenotypic plasticity. However, both the limits of plasticity and whether it will be sufficient to maintain local adaptation (or even lead to maladaptation) are less clear. Increased winter precipitation has been shown to lead to phenological delays and corresponding annual decreases in fitness in Columbian ground squirrels (Urocitellus columbianus). We took advantage of natural phenological variation (across elevations) in this species to better assess the extent of phenotypic plasticity in emergence dates and the relationships between emergence dates and individual annual fitness. We coupled a reciprocal translocation experiment with natural monitoring across two populations separated by â¼500 m in elevation. Individuals in both populations responded plastically to both spring temperature and winter precipitation. Translocated individuals adjusted their emergence dates to approach those of individuals in their adoptive populations but did differ significantly in their emergence dates from residents. There were no differences in annual fitness among treatment groups nor selection on emergence date within a year. Phenotypic plasticity is thus sufficient to allow individuals to respond to broad environmental gradients, but the influence of variation in emergence dates on annual fitness requires further investigation.
Assuntos
Adaptação Fisiológica/fisiologia , Hibernação/fisiologia , Sciuridae/fisiologia , Alberta , Altitude , Animais , Feminino , Gravidez , Chuva , Estações do Ano , TemperaturaRESUMO
The episodic production of large seed crops by some perennial plants (masting) is known to increase seed escape by alternately starving and swamping seed predators. These pulses of resources might also act as an agent of selection on the life histories of seed predators, which could indirectly enhance seed escape by inducing an evolutionary load on seed predator populations. We measured natural selection on litter size of female North American red squirrels (Tamiasciurus hudsonicus) across 28 years and five white spruce (Picea glauca) masting events. Observed litter sizes were similar to optimum litter sizes during nonmast years but were well below optimum litter sizes during mast years. Mast events therefore caused selection for larger litters ( ß'=0.25 ) and a lag load ( L=0.25 ) on red squirrels during mast years. Reduced juvenile recruitment associated with this lag load increased the number of spruce cones escaping squirrel predation. Although offspring and parents often experienced opposite environments with respect to the mast, we found no effect of environmental mismatches across generations on either offspring survival or population growth. Instead, squirrels plastically increased litter sizes in anticipation of mast events, which partially, although not completely, reduced the lag load resulting from this change in food availability. These results therefore suggest that in addition to ecological and behavioral effects on seed predators, mast seed production can further enhance seed escape by inducing maladaptation in seed predators through fluctuations in optimal trait values.
Assuntos
Sciuridae/fisiologia , Sementes , Adaptação Fisiológica , Animais , Feminino , Tamanho da Ninhada de Vivíparos/fisiologia , Picea/fisiologia , Dinâmica Populacional , Seleção Genética , YukonRESUMO
Evolutionary biologists have long trained their sights on adaptation, focusing on the power of natural selection to produce relative fitness advantages while often ignoring changes in absolute fitness. Ecologists generally have taken a different tack, focusing on changes in abundance and ranges that reflect absolute fitness while often ignoring relative fitness. Uniting these perspectives, we articulate various causes of relative and absolute maladaptation and review numerous examples of their occurrence. This review indicates that maladaptation is reasonably common from both perspectives, yet often in contrasting ways. That is, maladaptation can appear strong from a relative fitness perspective, yet populations can be growing in abundance. Conversely, resident individuals can appear locally adapted (relative to nonresident individuals) yet be declining in abundance. Understanding and interpreting these disconnects between relative and absolute maladaptation, as well as the cases of agreement, is increasingly critical in the face of accelerating human-mediated environmental change. We therefore present a framework for studying maladaptation, focusing in particular on the relationship between absolute and relative fitness, thereby drawing together evolutionary and ecological perspectives. The unification of these ecological and evolutionary perspectives has the potential to bring together previously disjunct research areas while addressing key conceptual issues and specific practical problems.
Assuntos
Adaptação Biológica , Evolução Biológica , Fenômenos Ecológicos e Ambientais , Aptidão Genética , Seleção GenéticaRESUMO
Organisms can affect one another's phenotypes when they socially interact. Indirect genetic effects occur when an individual's phenotype is affected by genes expressed in another individual. These heritable effects can enhance or reduce adaptive potential, thereby accelerating or reversing evolutionary change. Quantifying these social effects is therefore crucial for our understanding of evolution, yet estimates of indirect genetic effects in wild animals are limited to dyadic interactions. We estimated indirect phenotypic and genetic effects, and their covariance with direct effects, for the date of spring breeding in North American red squirrels (Tamiasciurus hudsonicus) living in an array of territories of varying spatial proximity. Additionally, we estimated indirect effects and the strength of selection at low and high population densities. Social effects of neighbours on the date of spring breeding were different from zero at high population densities but not at low population densities. Indirect phenotypic effects accounted for a larger amount of variation in the date of breeding than differences attributable to the among-individual variance, suggesting social interactions are important for determining breeding dates. The genetic component to these indirect effects was however not statistically significant. We therefore showcase a powerful and flexible method that will allow researchers working in organisms with a range of social systems to estimate indirect phenotypic and genetic effects, and demonstrate the degree to which social interactions can influence phenotypes, even in a solitary species.
Assuntos
Modelos Genéticos , Parto , Sciuridae/psicologia , Meio Social , Territorialidade , Animais , Feminino , Masculino , Sciuridae/genéticaRESUMO
Phenotypic plasticity-one individual's capacity for phenotypic variation under different environments-is critical for organisms facing fluctuating conditions within their lifetime. North American red squirrels (Tamiasciurus hudsonicus) experience drastic among-year fluctuations in conspecific density. This shapes juvenile competition over vacant territories and overwinter survival. To help young cope with competition at high densities, mothers can increase offspring growth rates via a glucocorticoid-mediated maternal effect. However, this effect is only adaptive under high densities, and faster growth often comes at a cost to longevity. While red squirrels can adjust hormones in response to fluctuating density, the degree to which mothers differ in glucocorticoid plasticity across changing densities remains unknown. Findings from our reaction norm approach revealed significant individual variation not only in a female red squirrel's mean endocrine phenotype but also in endocrine plasticity in response to changes in local density. Future work on proximate and ultimate drivers of variation in endocrine plasticity and maternal effects is needed, particularly in free-living animals experiencing fluctuating environments.