Early-life sibling conflict in Canada jays has lifetime fitness consequences.

While delaying natal dispersal can provide short-term benefits for juveniles, lifetime fitness consequences are rarely assessed. Furthermore, competition for limited positions on a natal territory could impose an indirect fitness cost on the winner if the outcome has negative effects on its siblings. We use radio-tracking and 58 years of nesting data in Ontario, Canada to examine the lifetime fitness consequences of sibling expulsion in the Canada jay (Perisoreus canadensis). Six weeks after fledging, intra-brood dominance struggles result in one 'dominant juvenile' (DJ) remaining on the natal territory after expelling its subordinate siblings, the 'ejectees' (EJs). Despite an older age-at-first-reproduction, DJs produced more recruits over their lifetime and had higher first-year survival than EJs, leading to substantially higher direct fitness. Even though DJs incurred an indirect fitness cost by expelling their siblings and there was no evidence that their presence on the natal territory increased their parents' reproductive output the following year, they still had substantially higher inclusive fitness than EJs. Our results demonstrate how early-life sibling conflict can have lifetime consequences and that such fitness differences in Canada jays are driven by the enhanced first-year survival of DJs pursuant to the early-summer expulsion of their sibling competitors.

Early-life experience shapes patterns of senescence in a food-caching passerine.

For many species, breeding performance increases through early adulthood followed by declines later in life. Although patterns of age-specific decline have been shown to vary between individuals, the factors that lead to this individual variation in the intensity of reproductive senescence are yet to be fully understood. We investigated whether early-life social status influenced age-related trends in the breeding performance of male Canada jays (Perisoreus canadensis), year-round residents of North America's boreal and sub-alpine forests. Shortly after young become nutritionally independent, intra-brood dominance struggles lead to one juvenile (Dominant Juvenile) remaining on the natal territory after expelling its subordinate siblings (Ejectees). First, we show via radio tracking that in our declining range-edge population Ejectees either join an unrelated pair (67%), form a breeding pair with another bird (28%) or occupy a territory alone (5%). Second, using 39 years of breeding data, we demonstrate that Ejectee males advanced laying dates and increased the annual number of nestlings until 6 years of age before declining, whereas Dominant Juvenile males advanced laying dates until 11 years and increased annual number of nestlings until 12 years of age before declining. This study documents clear variation in ageing patterns between dominant and expelled young, with implications for the role of early-life experiences and phenotypic quality in determining patterns of ageing.

Climate-driven carry-over effects negatively influence population growth rate in a food-caching boreal passerine.

Understanding how events throughout the annual cycle are linked is important for predicting variation in individual fitness, but whether and how carry-over effects scale up to influence population dynamics is poorly understood. Using 38 years of demographic data from Algonquin Provincial Park, Ontario, and a full annual cycle integrated population model, we examined the influence of environmental conditions and density on the population growth rate of Canada jays (Perisoreus canadensis), a resident boreal passerine that relies on perishable cached food for over-winter survival and late-winter breeding. Our results demonstrate that fall environmental variables, most notably the number of freeze-thaw events, carried over to influence late-winter fecundity, which, in turn, was the main vital rate driving population growth. These results are consistent with the hypothesis that warmer and more variable fall conditions accelerate the degradation of perishable stored food that is relied upon for successful reproduction. Future warming during the fall and winter may compromise the viability of cached food that requires consistent subzero temperatures for effective preservation, potentially exacerbating climate-driven carry-over effects that impact long-term population dynamics.

Fecal DNA metabarcoding helps characterize the Canada jay's diet and confirms its reliance on stored food for winter survival and breeding.

Accurately determining the diet of wild animals can be challenging if food items are small, visible only briefly, or rendered visually unidentifiable in the digestive system. In some food caching species, an additional challenge is determining whether consumed diet items have been previously stored or are fresh. The Canada jay (Perisoreus canadensis) is a generalist resident of North American boreal and subalpine forests with anatomical and behavioural adaptations allowing it to make thousands of arboreal food caches in summer and fall that are presumably responsible for its high winter survival and late winter/early spring breeding. We used DNA fecal metabarcoding to obtain novel information on nestling diets and compiled a dataset of 662 published and unpublished direct observations or stomach contents identifications of natural foods consumed by Canada jays throughout the year. We then used detailed natural history information to make informed decisions on whether each item identified to species in the diets of winter adults and nestlings was best characterized as 'likely cached', 'likely fresh' (i.e., was available as a non-cached item when it appeared in a jay's feces or stomach), or 'either possible'. Of the 87 food items consumed by adults in the winter, 39% were classified as 'likely cached' and 6% were deemed to be 'likely fresh'. For nestlings, 29% of 125 food items identified to species were 'likely cached' and 38% were 'likely fresh'. Our results support both the indispensability of cached food for Canada jay winter survival and previous suggestions that cached food is important for late winter/early spring breeding. Our work highlights the value of combining metabarcoding, stomach contents analysis, and direct observations to determine the cached vs. non-cached origins of consumed food items and the identity of food caches, some of which could be especially vulnerable to degradation through climate change.

Environmental conditions modulate compensatory effects of site dependence in a food-caching passerine.

Although density regulates the abundance of most wild animal populations by influencing vital rates, such as fecundity and survival, the mechanisms responsible for generating negative density dependence are unclear for many species. Site dependence occurs when there is preferential filling of high-quality territories, which results in higher per capita vital rates at low densities because a larger proportion of occupied territories are of high quality. Using 41 yr of territory occupancy and demographic data, we investigated whether site dependence was a mechanism acting to influence fecundity and, by extension, regulate a population of Canada Jays in Algonquin Provincial Park, Ontario, Canada. As predicted by site dependence, the proportion of occupied territories that were of high quality was negatively correlated with population density and periods of vacancy were shorter for high-quality territories than for low-quality territories. We also found evidence that per capita fecundity was positively related to the proportion of occupied territories that were of high quality, but only when environmental conditions, which influence the entire population, were otherwise poor for breeding. Our results suggest that site dependence likely plays a role in regulating this population but that environmental conditions can modulate the strength of density dependence.

Early-Life Corticosterone Body Condition Influence Social Status and Survival in a Food-Caching Passerine.

Individuals undergo profound changes throughout their early life as they grow and transition between life-history stages. As a result, the conditions that individuals experience during development can have both immediate and lasting effects on their physiology, behavior, and, ultimately, fitness. In a population of Canada jays in Algonquin Provincial Park, Ontario, Canada, we characterized the diet composition and physiological profile of young jays at three key time points during development (nestling, pre-fledge, and pre-dispersal) by quantifying stable-carbon (δ13C) and -nitrogen (δ15N) isotopes and corticosterone concentrations in feathers. We then investigated the downstream effects of early-life diet composition, feather corticosterone, and environmental conditions on a juvenile's social status, body condition, and probability of being observed in the fall following hatch. Across the three time points, the diet of Canada jay young was composed primarily of vertebrate tissue and human food with the proportion of these food items increasing as the jays neared dispersal. Feather corticosterone concentrations also shifted across the three time points, decreasing from nestling to pre-dispersal. Dominant juveniles had elevated corticosterone concentrations in their feathers grown pre-dispersal compared with subordinates. High body condition as nestlings was associated with high body condition as juveniles and an increased probability of being observed in the fall. Together, our results demonstrate that nestling physiology and body condition influence the social status and body condition once individuals are independent, with potential long-term consequences on survival and fitness.

The Effect of Egg Laying on Feather and Plasma Corticosterone Concentrations in Turkey (Meleagris gallopavo) Hens.

Phenological differences in energy demand (i.e., reproductive status) might influence the measurement of corticosterone. The objective of this study was to compare corticosterone concentrations in feathers (FCORT) and plasma (PCORT) for turkey hens before and during egg laying. Secondary feathers 1 and 3, and a plasma sample were collected from 50 hens at 30 weeks (before egg laying) from two purebred lines. The hens were reexamined during lay (45 weeks) to collect regrown feathers and plasma samples. Corticosterone concentrations were measured using an ELISA. Linear mixed models were used to assess the effect of genetic line (A vs. B) and period (pre-lay vs. lay) on FCORT and PCORT levels. An increase in FCORT during lay was detected for line B (p < 0.0001), but not line A (p = 0.3076). An increase in FCORT during lay was detectable in both feather types, but there was a difference between secondary 1 and 3 in FCORT concentration within each line studied. Conversely, PCORT decreased between the pre-lay and lay periods for both lines, although the decrease was more substantial for line A (p < 0.0001). Differences in metabolic investment in egg production between the two genetic lines may explain the different FCORT response during lay. The results from this study provide insight into how periods of high energy demand may influence corticosterone which should be considered when interpreting results.

Raising young with limited resources: supplementation improves body condition and advances fledging of Canada Jays.

Food availability early in life can play a vital role in an individual's development and success, but experimental evidence for the direct effects of food on body condition, physiology, and survival of young animals in the wild is still relatively scarce. Food-caching Canada Jays (Perisoreus canadensis) begin breeding in the late winter and, therefore, rely on either cached food or seemingly limited quantities of fresh food to feed nestlings in the early spring. Using a 2-yr food supplementation experiment conducted during the nestling period and 40 yr of observational data on food supplemented by the public, we examined whether food quantity during early life influenced the physiology, body condition, timing of fledging, and survival of young Canada Jays in Algonquin Provincial Park, Ontario, Canada. Experimental food supplementation of breeding pairs advanced the fledging date of young by 24% (5.5 d) compared to controls. In 1 yr of the experiment, nestlings raised on experimentally supplemented territories had lower feather corticosterone concentrations and were in higher body condition than controls. Across treatment and control nests, young that successfully fledged had lower concentrations of feather corticosterone and were in higher body condition than those that did not fledge. Based on 40 yr of observational data, nestling body condition was positively related to the degree of food supplementation by park visitors and nestlings in higher body condition were more likely to be observed in the population in the following fall. Our results demonstrate how food availability early in life can have important downstream consequences on metrics related to individual fitness, including first year survival.

Research Note: Quantifying corticosterone in turkey (Meleagris gallopavo) feathers using ELISA.

The measurement of corticosterone in feathers is an appealing tool for assessing glucocorticoids in wild and domestic bird species. Feather corticosterone measurements can be performed noninvasively and can provide a means for comparing glucocorticoid secretion between individual birds; thus, such measurements can be used to assess the welfare of domestic poultry. The focus of this study was to assess the validity of detecting corticosterone in turkey (Meleagris gallopavo) feathers using an ELISA and investigate differences between genetic lines. Primary feather 9 was obtained at a processing plant from 123 individuals from 3 different purebred turkey lines (line A [N = 46], line B [N = 24], and line C [N = 53]). Assay validation tests were performed using a commercially available ELISA kit (Cayman Chemicals, Cedarlane Labs, Canada). Indicators of accuracy, recovery, precision, and sensitivity were sufficient. Significant differences in feather corticosterone concentration between the 3 lines were observed. Line C had significantly higher feather corticosterone than line A (P < 0.0001) and line B (P = 0.036). These results indicate that the quantification of feather corticosterone using an ELISA is a valid method for assessing glucocorticoid levels in turkeys. This is the first report of differences in feather corticosterone between different purebred turkey lines. Differences observed between purebred lines provide an intriguing basis for further investigation into the genetic parameters of glucocorticoid levels in turkeys.

Autumn freeze-thaw events carry over to depress late-winter reproductive performance in Canada jays.

Evidence suggests that range-edge populations are highly vulnerable to the impacts of climate change, but few studies have examined the specific mechanisms that are driving observed declines. Species that store perishable food for extended periods of time may be particularly susceptible to environmental change because shifts in climatic conditions could accelerate the natural degradation of their cached food. Here, we use 40 years of breeding data from a marked population of Canada jays (Perisoreus canadensis) located at the southern edge of their range in Algonquin Provincial Park, Ontario, to examine whether climatic conditions prior to breeding carry over to influence reproductive performance. We found that multiple measures of Canada jay reproductive performance (brood size, nest success and nestling condition) in the late winter were negatively correlated with the number of freeze-thaw events the previous autumn. Our results suggest that freeze-thaw events have a significant detrimental impact on the quality and/or quantity of cached food available to Canada jays. Future increases in such events, caused by climate change, could pose a serious threat to Canada jays and other food caching species that store perishable foods for long periods of time.

Length polymorphisms at two candidate genes explain variation of migratory behaviors in blackpoll warblers (Setophaga striata).

Migratory behaviors such as the timing and duration of migration are genetically inherited and can be under strong natural selection, yet we still know very little about the specific genes or molecular pathways that control these behaviors. Studies in candidate genes Clock and Adcyap1 have revealed that both of these loci can be significantly correlated with migratory behaviors in birds, though observed relationships appear to vary across species. We investigated geographic genetic structure of Clock and Adcyap1 in four populations of blackpoll warblers (Setophaga striata), a Neotropical-Nearctic migrant that exhibits geographic variation in migratory timing and duration across its boreal breeding distribution. Further, we used data on migratory timing and duration, obtained from light-level geolocator trackers to investigate candidate genotype-phenotype relationships at the individual level. While we found no geographic structure in either candidate gene, we did find evidence that candidate gene lengths are correlated with five of the six migratory traits. Maximum Clock allele length was significantly and negatively associated with spring arrival date. Minimum Adcyap1 allele length was significantly and negatively associated with spring departure date and positively associated with fall arrival date at the wintering grounds. Additionally, we found a significant interaction between Clock and Adcyap1 allele lengths on both spring and fall migratory duration. Adcyap1 heterozygotes also had significantly shorter migration duration in both spring and fall compared to homozygotes. Our results support the growing body of evidence that Clock and Adcyap1 allele lengths are correlated with migratory behaviors in birds.

Quantifying corticosterone in feathers: validations for an emerging technique.

Feather corticosterone measurement is becoming a widespread tool for assessing avian physiology. Corticosterone is deposited into feathers during growth and provides integrative and retrospective measures of an individual's hypothalamic-pituitary-adrenal (HPA) axis function. Although researchers across disciplines have been measuring feather corticosterone for the past decade, there are still many issues with the extraction and measurement of corticosterone from feathers. In this paper, we provide several directives for refining the methodology for feather hormone analysis. We compare parallelism between the standard curve and serially diluted feather tissue from wild turkeys, Canada jays, and black-capped chickadees to demonstrate the wide applicability across species. Through a series of validations, we compare methods for feather preparation, sample filtration and extract reconstitution prior to corticosterone quantification using a radioimmunoassay. Higher corticosterone yields were achieved following pulverization of the feather however, more variation between replicates was observed. Removal of the rachis also increased the amount of corticosterone detected per unit mass while glass versus paper filters had no effect, and using ethanol in the reconstution buffer decreased intra-assay variation. With these findings and continued methodological refinement, feather corticosterone has the potential to be a powerful tool for both ecologists and physiologists working with historical and contemporary specimens.

The effect of elbow flexor fatigue on spine kinematics and muscle activation in response to sudden loading at the hands.

Sudden loads, originating at either the hands or the feet, can cause injury to spine structures. As muscles are primarily responsible for stabilization following a perturbation, the effect of spine muscle fatigue in this context has been well investigated. However, the effect of fatigue of arm muscles, which can help control perturbations originating at the hands, on the spine is unknown. The purpose of this study was to determine if the magnitude of spine flexion or the pre-activation, reflex amplitude, and reflex latency of spine muscles were altered by elbow flexor fatigue during a sudden loading (6.8 kg) perturbation at the hands. Elbow flexor fatigue was induced by an isometric 30% maximal elbow flexion moment until failure. Results demonstrate that spine kinematics were not altered in the presence of elbow flexor fatigue. Small magnitude differences in trunk muscle pre- and peak activation indicate that the presence of elbow flexor fatigue does not necessitate substantially greater spine muscle action under the tested conditions. Despite fatigued elbow flexors, the arm muscles were sufficiently able to control the perturbation. Interestingly, 5/14 participants demonstrated altered reflex latencies in all observed muscles that lasted up to 10 min after the fatiguing task.