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There is mounting evidence from many species that physiological state can be reflected in external cues, such as the structure of vocalizations. North American red squirrels ( Tamiasciurus hudsonicus (Erxleben, 1777)) produce individually distinct territorial vocalizations, called rattles, containing information relating to both a proximate stressor and reproductive condition. Given the importance of these vocalizations in communicating territory ownership, and their ability to contain cues relating to physiological state, we hypothesized that information regarding individual circulating glucocorticoid concentrations is transmitted between conspecifics within an audible range. We analysed fecal cortisol metabolite (FCM) concentrations from 129 red squirrels, comparing focal individual FCMs to conspecific FCMs within an acoustic neighbourhood. We found support for our hypothesis through a significant positive relationship between the FCM concentrations of focal squirrels and their neighbours independent of local density. These findings demonstrate that individual physiology interacts with conspecific physiology at ecologically relevant spatial scales and suggest that the social relationships between individual red squirrels may play a larger role in dictating individual responses to the environment. 

Despite considerable research on reproductive timing, factors influencing intraspecific variation in gestation length in wild mammals have received limited attention, largely due to a lack of high-resolution data. We examined potential drivers of gestation length variation in a wild population of North American red squirrels ( Tamiasciurus hudsonicus Erxleben, 1777) in Yukon, Canada. Drawing on 27 years of conception and parturition data, we found that gestation length was highly conserved, with minimal variation (35.4 ± 1.49 days; mean ± standard deviation). Gestation length was unaffected by maternal age, litter size, or litter sex ratio. Likewise, the anticipation of an upcoming food pulse, caused by synchronous conifer seed masting, had no statistically significant influence on gestation length. This finding stands in contrast to other reproductive traits in red squirrels, which are known to shift in response to food pulses. Overall, our results suggest that gestation length in red squirrels is buffered against environmental variability, likely due to stabilizing selection or strong heritability. 

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. 

Abstract Female advertisement of reproductive state and receptivity has the potential to play a large role in the mating systems of many taxa, but investigations of this phenomenon are underrepresented in the literature. North American red squirrels (Tamiasciurus hudsonicus) are highly territorial and engage in scramble competition mating, with males converging from spatially disparate territories to engage in mating chases. Given the narrow estrus window exhibited in this species, the ubiquitous use of vocalizations to advertise territory ownership, and the high synchronicity of males arriving from distant territories, we hypothesized that female vocalizations contain cues relating to their estrous state. To test this hypothesis, we examined the spectral and temporal properties of female territorial rattle vocalizations collected from females of known reproductive condition over 3 years. While we found no distinct changes associated with estrus specifically, we did identify significant changes in the spectral characteristics of rattles relating to both female body mass and reproductive state relative to parturition. To the best of our knowledge, this is the first evidence of changes in vocal characteristics associated with late pregnancy in a nonhuman mammal. 

In fluctuating environments, phenotypic mismatching may reflect an error-avoidance bias that enhances a mother squirrel’s overall lifetime fitness. 

ABSTRACT Microbial transmission is hypothesised to be a major benefit of sociality, facilitated by affiliative behaviours such as grooming and communal nesting in group‐living animals. Whether microbial transmission is also present in animals that do not form groups because territoriality limits interactions and prevents group formation remains unknown. Here, we investigate relationships among gut microbiota, population density and dynamic behavioural and spatial measures of territoriality in wild North American red squirrels (Tamiasciurus hudsonicus). Periods of high population density predicted population‐level gut microbial homogeneity but individual‐level diversification, alongside changes in obligately anaerobic, non‐sporulating taxa indicative of social transmission. Microbial alpha‐diversity increased with more frequent territorial intrusions, and pairs with stronger intrusion‐based social associations had more similar gut microbiota. As some of the first evidence for social microbial transmission in a solitary system, our findings suggest that fluctuations in density and territorial behaviours can homogenise and diversify host microbiomes among otherwise non‐interacting animals. 

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.