Tamiasciurus Hudsonicus Research Articles

Boom and bust: the effects of masting on seed predator range dynamics and trophic cascades

AbstractAimSpatiotemporal variation in resource availability is a strong driver of animal distributions. In the northern hardwood and boreal forests of the northeastern United States, tree mast events provide resource pulses that drive the population dynamics of small mammals, including the American red squirrel (Tamiasciurus hudsonicus), a primary songbird nest predator. This study sought to determine whether mast availability ameliorates their abiotic limits, enabling red squirrel elevational distributions to temporarily expand and negatively impact high‐elevation songbirds.LocationNortheastern United States.MethodsWe used two independent datasets to evaluate our hypotheses. First, we fit a dynamic occupancy model using data from camera trap surveys to evaluate red squirrel distributional responses to pulses in the tree mast. We also assessed population responses using systematic auditory surveys analysed with an open‐population binomial mixture model. Further, we used modelled red squirrel abundance in nest‐survival models to evaluate whether their abundance is correlated with the daily nest survival of three songbird species.ResultsThe tree mast provided a critical resource pulse that resulted in a two‐fold increase in the annual elevational distribution of red squirrels. The elevational distribution of red squirrels ranged from a minimum of ~450 m (range: 663–1145 m asl) following two consecutive years without a masting event to a maximum of over 1000 m (range: 443–1545 m asl) after a large mast event. The daily nest survival of three songbird species tended to decline with an increase in the abundance of red squirrels.Main ConclusionsTree mast is a central biological phenomenon in many temperate and boreal forests. This study reveals how this resource pulse results in range changes in a small mammal that is both a seed and bird predator, as well as prey for many carnivores. Thus, understanding this phenomenon can inform the conservation and management of northern forests, including breeding songbirds.

A future food boom rescues the negative effects of early-life adversity on adult lifespan in a small mammal.

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.

Motion‐sensitive cameras track population abundance changes in a boreal mammal community in southwestern Yukon, Canada

AbstractMotion‐sensitive cameras are commonly used to monitor wildlife occupancy rates; however, few studies have assessed whether data from cameras are correlated with density estimates obtained from more traditional labor‐intensive methods such as those based on capture‐mark‐recapture. We used data from a boreal forest community to test whether camera data were correlated with densities estimated from independent monitoring methods. We placed 72 covert cameras in the forest around Lhù'ààn Mân' (Kluane Lake), Yukon, Canada, for 7 years and tracked changes in population densities by camera hit rates. We independently estimated population densities of snowshoe hares (Lepus americanus) and red squirrels (Tamiasciurus hudsonicus) using capture‐mark‐recapture via live trapping, and Canada lynx (Lynx canadensis), coyotes (Canis latrans), and moose (Alces americanus) by snow track transects. Density estimates obtained from conventional aerial surveys were also periodically available for moose. Except for red squirrels, camera hit rates were highly correlated with population density estimates obtained by traditional methods, including across a large range of estimated densities corresponding to cyclic population dynamics in several species. Accordingly, we infer that motion‐sensitive cameras could supplement or replace traditional methods for monitoring key species in boreal forest food webs. Using cameras to monitor population change has several advantages; they require less effort in the field, are non‐invasive compared to live‐trapping, include multiple species at the same time, and rely less on weather than either aerial surveys or snow track transects. Tracking changes across the vast boreal forest is becoming increasingly necessary because of climate and landscape change and our data validate the use of motion‐sensitive cameras to provide a useful quantitative method for state‐of‐the‐environment reporting.

Reproductive state alters vocal characteristics of female North American red squirrels (Tamiasciurus hudsonicus)

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.

Inferring condition in wild mammals: body condition indices confer no benefit over measuring body mass across ecological contexts.

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.

Consistent spatial patterns in microbial taxa of red squirrel gut microbiomes.

Gut microbiomes are diverse ecosystems whose drivers of variation remain largely unknown, especially in time and space. We analysed a dataset with over 900 red squirrel (Tamiasciurus hudsonicus) gut microbiome samples to identify the drivers of gut microbiome composition in this territorial rodent. The large-scale spatiotemporal replication in the data analysed was an essential component of understanding the assembly of these microbial communities. We identified that the spatial location of the sampled squirrels in their local environment is a key contributor to gut microbial community composition. The non-core gut microbiome (present in less than 75% of gut microbiome samples) had highly localised spatial patterns throughout different seasons and different study areas in the host squirrel population. The core gut microbiome, on the other hand, showed some spatial patterns, though fewer than in the non-core gut microbiome. Environmental transmission of microbiota is the likely contributor to the spatiotemporal distribution observed in the North American red squirrel gut microbiome.

Experimental manipulation of perceived predation risk alters survival, cause of death, and demographic patterns in juvenile snowshoe hares

Perceived predation risk alters prey behaviour and physiology, but few studies have examined downstream consequences on prey demography in wild populations. Perceived predation risk could alter adult reproductive performance via reduced investment in offspring quality and post-birth care. We manipulated perceived predation risk in snowshoe hare ( Lepus americanus Erxleben, 1777) by exposing pregnant mothers to chases by a domestic dog ( Canis familiaris Linnaeus, 1758). Litter size was comparable between risk-augmented and control groups, but treated females had more stillbirths and gave birth to leverets of lower body condition. Leverets from risk-augmented females had 88% higher 40-day mortality rate. Maternally preventable causes of death like starvation or predation by red squirrel ( Tamiasciurus hudsonicus (Erxleben, 1777)) caused this difference, particularly during the nursing period. Risk-augmented mothers were always more active than controls, but the difference was greatest during the nursing period. We found that perceived predation risk reduces maternal productivity pre- and post-partum, implying downstream consequences to populations. Because our treatment ended before parturition, we can link offspring performance such as survival and behaviour specifically to maternal life-history trade-offs, which has not been shown in a wild mammal.

Long-term monitoring in the boreal forest reveals high spatio-temporal variability among primary ecosystem constituents

The boreal forest, the world’s largest terrestrial biome, is undergoing dramatic changes owing to anthropogenic stressors, including those of climate change. To track terrestrial ecosystem changes through space and time, robust monitoring programs are needed that survey a variety of ecosystem constituents. We monitored white spruce (Picea glauca) cone crops, berry (Empetrum nigrum, Shepherdia canadensis) production, above-ground mushroom abundance, and the abundance of small mammals (Clethrionomys rutilus,Peromyscus maniculatus), North American red squirrels (Tamiascirus hudsonicus), snowshoe hares (Lepus americanus), and carnivores (Lynx canadensis,Canis latrans,Vulpes vulpes,Martes americana,Mustela erminea) across 5 sites in the Yukon, Canada. Monitoring began in 1973 at Lhù’ààn Mân’ (Kluane Lake) and additional protocols were added until a complete sequence was fixed in 2005 at all 5 sites and continued until 2022. White spruce cone counts show mast years at 3–7-year intervals. Ground berries and soapberry counts were highly variable among sites and counts did not correlate among sites or between years for different species. Red-backed voles showed clear 3–4-year cycles at Kluane and probably at the Mayo and Watson Lake sites, but showed only annual cycles in Whitehorse and Faro. Snowshoe hares fluctuated in 9–10-year cycles in a travelling wave, peaking one year earlier at Watson Lake but in synchrony at all other sites, with no clear sign of peak density changing or cyclic attenuation over the last 50 years. Red squirrel numbers at Kluane exhibit marked inter-year variability, driven mainly by episodic white spruce cone crops and predation from Canada lynx and coyotes as hare densities undergo cyclic decline. Snow track counts to index mammalian predators have been conducted on our Kluane and Mayo sites, indicating that lynx numbers rise and fall with a 1–2-year lag at these two sites, tracking the hare cycle. Coyotes and lynx at Kluane peak together following the hare cycle, but coyote counts are also depressed during deep snow years. To summarize, we noted considerable inter-site variability in the population dynamics of many boreal forest ecosystem constituents, but the keystone species (snowshoe hare, Canada lynx) exhibit remarkably similar population trends across the region. We continue to monitor wildlife abundance, cone crops, berry production, and mushroom biomass to determine changes associated with increasing temperature and fluctuating rainfall. The Yukon boreal forest is changing as climate shifts, but the changes are slow, variable across sites, taxa specific, and of uncertain predictability.

Divergent or convergent: how do forest carnivores use time in the Greater Yellowstone Ecosystem?

Abstract Divergent activity can change the intensity of species interactions, largely affecting species distributions and abundances, and consequently influencing the composition and function of ecological communities. Few assessments of activity patterns have focused questions around different resource constraints or have examined varying time frames when interaction strengths are expected to increase. We evaluated how activity among carnivores and their prey shifted from early to late winter, coinciding with a presumed decrease in food resources for carnivores, and we measured time between species detections within a camera station. Our study species were three forest carnivores—Pacific martens (Martes caurina), Rocky Mountain red foxes (Vulpes vulpes macroura), coyotes (Canis latrans); and two of their prey—American red squirrels (Tamiasciurus hudsonicus), and snowshoe hares (Lepus americanus). We sampled these species across an extensive network of cameras (n = 107) during the 2014–2017 winter seasons in the Greater Yellowstone Ecosystem, Wyoming. We generated kernel density plots for timing of photographs and calculated the coefficient of overlap among density plots for our predators and prey during early and late winter. Furthermore, we calculated the time-between-detections (i.e., hours) among forest carnivores. We found no consistent trends in time-between-detections across our species pairs. Pacific martens exhibited cathemeral activity that aligned with the peaks in activity of the two prey species. Temporal overlap between coyote and red fox activities was small in early winter, whereas coyotes modified activity in late winter such that they more closely aligned with red foxes. This intraguild convergence of activity may reflect an increase in resource constraints and have consequences for competitive interactions between these two canids. Our study supports the notion that variation in time is an important axis in facilitating coexistence among these forest carnivores and prey species.

Differential impacts of spruce beetle outbreaks on snowshoe hares and red squirrels in the southern Rocky Mountains

Spruce beetles (Dendroctonus rufipennis) have impacted millions of acres of Engelmann spruce (Picea engelmannii) – subalpine fir (Abies lasiocarpa) forest in North America over the past decade, resulting in the most extensive outbreak in recorded history. This dramatic alteration of forest composition and structure has precipitated numerous changes to forest ecology and ecosystem services. Among the least studied of these changes are impacts to wild mammals, including snowshoe hares (Lepus americanus) and red squirrels (Tamiasciurus hudsonicus). We sampled a chronosequence of spruce-fir stands along a gradient of ‘years elapsed since spruce beetle outbreak’ (YSO) in order to estimate impacts to abundance of these two species in the southern Rocky Mountains. Snowshoe hare abundance was not related to YSO, at least in the first decade post-outbreak. Instead, hare abundance during this period was positively related to horizontal cover, especially that due to stem density of small diameter subalpine fir. Notably, snowshoe hare abundance was negatively related to stem density of small diameter Engelmann spruce, suggesting that elements of horizontal cover may not be uniformly beneficial to hares. Hare abundance was also negatively related to ground cover, which could help explain the lack of relationship to YSO, assuming reduction in overstory canopy would lead to increases in ground cover. Red squirrel abundance was negatively related to YSO and outbreak severity (i.e., basal area of large diameter dead trees). This was likely due to diminished cone crops in impacted areas, which red squirrels cache and rely on heavily to sustain them through the winter. Basal area of remaining large live fir trees was not related to squirrel abundance, suggesting that regeneration of spruce and associated cone crops may be necessary for recovery of red squirrels, which may take several decades.

Widespread, symmetric, noninflammatory alopecia associated with follicular dysplasia in the American red squirrel (Tamiasciurus hudsonicus).

This report describes a unique pattern of alopecia in 8 American red squirrels (Tamiasciurus hudsonicus) from 2013 to 2021. All animals were juveniles; 6 were female and 2 were male. Seven presented between September and November, and one presented in April. All squirrels had widespread, bilaterally symmetric, noninflammatory, well-demarcated alopecia involving the entire trunk and legs and normal hair on their muzzle and dorsal surfaces of their paws. Six months later, a normal hair coat had grown on 2 of the animals, which were littermates. Hair fully grew 2 months later in another animal. Histopathology of the alopecic skin was performed in 7 of 8 animals. The following changes were noted: bent and coiled hairs, perforating folliculitis, melanin clumping, and distortion of hair shafts. Based on features of follicular dysplasia and apparent seasonality, this condition has some similarities to canine seasonal flank alopecia. A genetic etiology is suspected.

Survey of Toxoplasma gondii in Urban and Rural Squirrels (Sciuridae) in Manitoba, Canada.

The coccidian parasite Toxoplasma gondii is found worldwide infecting warm-blooded vertebrates. Felids are the definitive hosts; other species act as intermediate hosts. Squirrels (Sciuridae) generally have high population densities in cities and forage and cache food on the ground, where they may come into contact with T. gondii oocysts or be preyed upon by cats and other carnivores. This environment might make squirrels important intermediate hosts of T. gondii in cities, and infection rates could indicate environmental levels of oocysts in soil. We investigated whether urban squirrels would be more exposed to T. gondii infection than rural squirrels with samples collected from American red squirrels (Tamiasciurus hudsonicus), eastern grey squirrels (Sciurus carolinensis), northern flying squirrels (Glaucomys sabrinus), and least chipmunks (Tamias minimus) in and around Winnipeg, Manitoba, Canada. We tested 230 tissue samples from 46 squirrels for T. gondii DNA by quantitative PCR and 13 serum samples from grey squirrels for T. gondii antibodies by competitive ELISA. We found no evidence of infection in any squirrel, indicating that squirrels are probably not important intermediate hosts of T. gondii in cities and that consumption of oocysts in the soil in general may not be an important contributor to transmission in colder environments.

A NEW SPECIES OF FLEA (SIPHONAPTERA: CTENOPHTHALMIDAE) PARASITIZING VOLES AT HIGH ELEVATIONS IN THE GREAT SMOKY MOUNTAINS OF TENNESSEE AND NORTH CAROLINA.

Both sexes of Catallagia appalachiensis n. sp. are described from high elevation spruce-fir forests in Sevier County, Tennessee and adjoining Swain County, North Carolina in the Great Smoky Mountains National Park. The type host of the new flea is the southern red-backed vole, Myodes gapperi (Vigors) (25 flea specimens), although small numbers of specimens were also collected from a sympatric northern short-tailed shrew, Blarina brevicauda (Say) (2 fleas), a red squirrel, Tamiasciurus hudsonicus (Erxleben) (1 flea), and a North American deer mouse, Peromyscus maniculatus (Wagner) (1 flea). Infestation prevalences for these hosts are provided. The new species is compared morphologically with other known species of Catallagia, in particular with Catallagia borealis, the only other described congeneric flea in eastern North America. This is the first new species of flea to be described from the eastern United States since 1980.

Beaver activity and red squirrel presence predict bird assemblages in boreal Canada

AbstractWetlands and predation in boreal ecosystems play essential roles throughout the breeding season for bird assemblages. We found a positive association of beaver activity and a negative influence of American red squirrels (Tamiasciurus hudsonicus) on bird assemblages. We used a multispecies hierarchical model to investigate whether bird communities differ between two major wetland habitats in boreal Canada: beaver ponds and peatland ponds. In addition to including variables such as forest cover and latitude, we adopted a structural equation model approach to estimate the occupancy of American red squirrels and its potential influence on bird communities. Using automated recording stations deployed at 50 ponds, we detected 96 bird species in 2018 and 2019. Bird species were grouped into four taxonomic guilds according to their habitat successional requirements: early successional species, late successional species, generalists, and wetland species. Beaver ponds harbored higher species richness, a pattern driven primarily by early successional species. The occupancy of almost a quarter of the species was lower in the presence of red squirrels. Late successional species responded positively to the cover of forest surrounding the pond. Our results highlight the value of considering acoustic data of red squirrels to quantify habitat quality in boreal forests. We conclude that beaver activity shapes bird assemblages through modification of their habitat, and that some bird guilds are associated negatively with the presence of American red squirrels.

Benefits of living closer to kin vary by genealogical relationship in a territorial mammal.

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.

Change in altitudinal distribution of Newfoundland Gray-cheeked Thrush ( Catharus minimus minimus ) revealed through historical stop-level breeding bird survey data

Breeding bird survey (BBS) data are typically analyzed at the route level in which the average or combined total number of individuals across the 50 stops is treated as a single observation. However, analysis at the stop level may provide information on finer-scale habitat affiliations of species within the broader landscape, as well as changes in species distribution over time. Though once abundant, the Newfoundland Gray-cheeked Thrush (<em>Catharus minimus minimus</em>) underwent a dramatic population collapse across most of the island during the 1980s and 1990s and today is most common in montane forests. The reason for this collapse is not well understood but introduced North American red squirrels (<em>Tamiasciurus hudsonicus</em>), which colonized most forested areas on the island below ~400 m from the late 1970s through the mid-1990s, have been identified as a possible cause. To improve our understanding of changes in the elevational distribution of Gray-cheeked Thrushes, we compared stop-level BBS data from two periods. Our historic period from 1974-1984 corresponds with the initial expansion phase of introduced red squirrels, whereas that from 2000-2015 reflects a period during which the colonization of suitable habitat by squirrels was complete. We used 17 routes (850 stops) where the thrushes had been observed and there were at least 3 surveys completed during each period. Surveys during the 1970s and 1980s indicated that Gray-cheeked Thrushes were abundant at all elevations sampled by the BBS (0-450 m), with higher incidence below 100 m and above 300 m; in contrast, recent surveys indicate that the species is now essentially absent below 350 m. This finding highlights the value of historical stop-level BBS data for reconstructing changes in species distributions and is consistent with the hypothesis that red squirrels have adversely affected Newfoundland Gray-cheeked Thrushes.

Density-dependent plasticity in territoriality revealed using social network analysis.

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.

Weather and nest cavity characteristics influence fecundity in mountain chickadees.

Examining direct and indirect effects on reproduction at multiple scales allows for a broad understanding of species' resilience to environmental change. We examine how the fecundity of the mountain chickadee (Poecile gambeli), a secondary cavity-nesting, insectivorous bird, varied in relation to factors at three scales: regional weather conditions, regional- and site-level food availability, site-level community dynamics, and nest-level cavity characteristics. We hypothesized that earlier laying dates and higher fecundity (clutch size, nest survival, brood size) would be associated with milder climatic conditions, increased food from insect outbreaks, lower densities of conspecifics and nest predators (red squirrel; Tamiasciurus hudsonicus), and safer (smaller, higher) cavities. We collected data on laying date, clutch size, brood size, nest fate (success/failure), and cavity characteristics from 513 mountain chickadee nests in tree cavities in temperate mixed coniferous-broadleaf forest in interior British Columbia, Canada, from 2000 to 2011. We surveyed annual abundances of mountain chickadees and squirrels using repeated point counts, and mountain pine beetle (Dendroctonus ponderosae) and lepidopteran defoliators by monitoring host trees and by using regional-scale aerial overview forest insect survey data. We used weather data (temperature, rain, snow) from a local Environment and Climate Change Canada weather station. We modeled laying date, clutch size, daily nest survival, and brood size as a function of predictors at regional-, site-, and nest-scales. Measures of fecundity varied dramatically across years and spatial scales. At the regional (study-wide) scale, chickadees laid earlier and larger first clutches in warmer springs with minimal storms, and daily nest survival (DSR) increased with a 2-year lag in growing season temperature. Despite a doubling of mountain chickadee density that roughly accompanied the outbreaks of mountain pine beetle and lepidopteran defoliators, we found little evidence at the site scale that fecundity was influenced by insect availability, conspecific density, or predator density. At the nest scale, DSR and brood size increased with clutch size but DSR declined with nest cavity size indicating a positive reproductive effect of small-bodied cavity excavators. Double-brooding, rare in chickadees, occurred frequently in 2005 and 2007, coinciding with early breeding, high food availability from insect outbreaks, and warm spring temperatures with 0-1 spring storms. Our results support the idea that fecundity in secondary cavity-nesting species is impacted directly and indirectly by weather, and indirectly through changes in community dynamics (via cavity resource supply). We stress the importance of adopting holistic, community-level study frameworks to refine our understanding of fecundity in opportunistic and climate-sensitive species in future.

The glucocorticoid response to environmental change is not specific to agents of natural selection in wild red squirrels

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.

Pulsed resources and the resource‐prediction strategy: a field‐test using a 36‐year study of small mammals

Pulsed resource environments are known for their marked variations in resource availability over space and time. Animals living in such environments usually increase reproduction after resources become available. Some small mammal populations, however, may use environmental cues that precede large crops of seeds (e.g. pollen, flowers, unripe seeds) to ‘predict' mast‐seeding events and reproduce in advance. In other words, these populations exhibit a resource‐prediction strategy based on future fitness prospects. Habitat selection might play a key role in the resource‐prediction strategy, but our understanding of habitat selection in pulsed resource environments is extremely limited. We used a 36‐year dataset on small mammal trapping and seed availability collected in Maine (USA) to test if American red squirrels Tamiasciurus hudsonicus, eastern chipmunks Tamias striatus, white‐footed mice Peromyscus leucopus and southern red‐backed voles Myodes gapperi select habitats based on upcoming resources and examine the consequences to individual fitness. Small mammal trapping was conducted in summer before seeds were available, thus trapping occurred when only the cues of an upcoming mast event were available. Using home‐range estimations and capture data, we performed resource selection analyses to assess if individuals select habitat to maximize future resource acquisitions. Using capture–mark–recapture models, we also estimated individual survival (a proxy for fitness). Our results do not support habitat selection as a mechanism underlying the resource‐prediction hypothesis, indicating that individuals do not select habitats based on the upcoming seed availability. Nevertheless, we found that white‐footed mice with home ranges in areas characterized by a greater quantity of white oak acorns have up to a 15% higher survival rate. Our empirical study advances the understanding of predator response to pulsed resources by not supporting a key mechanism thought to underpin the anticipatory responses that have been observed in many ecosystems worldwide.