Perisoreus Canadensis Research Articles

Learning about Extraordinary Beings: Native Stories and Real Birds

Oral traditions of Indigenous American peoples (as well as those of other Indigenous peoples) have long been discussed with regard to their reliability as metaphorical accounts based upon historical knowledge. I explore this debate using stories to discuss the importance of the role of Corvidae in Indigenous knowledge traditions and how these stories convey information about important socioecological relationships. Contemporary science reveals that Corvids important in cultural traditions were companions to humans and important components of the ecology of the places where these peoples lived. Ravens, Crows, Jays, and Magpies are identified as having special roles as cooperators, agents of change, trickster figures, and important teachers. Canada (or Gray) Jays serve as trickster/Creator of the Woodland Cree people, Wisakyjak. Magpies won the Great Race around the Black Hills to determine whether humans would eat bison or vice versa. I analyze these stories in terms of their ecological meaning, in an effort to illustrate how the stories employ dramatic settings to encourage respect and fix relationships in the sociocultural memory of the people.

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 41yr 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.

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 40yr 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 1yr 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 40yr 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.

New avian breeding records for Kugluktuk, Nunavut

New breeding records for 10 species of tundra and boreal nesting birds were documented near the community of Kugluktuk (Nunavut, Canada) over the course of the 2015 and 2016 breeding seasons and incidentally in 2017 and 2018. These species include American Wigeon (Mareca americana), Mallard (Anas platyrhynchos), Northern Shoveler (Anas clypeata), Green-winged Teal (Anas carolinensis), Greater Scaup (Aythya marila), Say’s Phoebe (Sayornis saya), Canada Jay (Perisoreus canadensis), Greycheeked Thrush (Catharus minimus), Yellow-rumped Warbler (Setophaga coronata), and Dark-eyed Junco (Junco hyemalis). Previously unpublished breeding evidence for Bald Eagle (Haliaeetus leucocephalus) is also discussed along with suspected breeding of Barn Swallow (Hirundo rustica), Bohemian Waxwing (Bombycilla garrulus), White-throated Sparrow (Zonotrichia albicollis), and Pine Grosbeak (Pinicola enucleator). These records represent the first described breeding occurrences for these species in the Kitikmeot region, or new records for the mainland of Nunavut. A lack of historical ornithological survey effort in this area has likely led to the diversity of these previously unrecorded breeding observations. These results highlight the need to increase geographic coverage of bird surveys in northern Canada to more accurately delineate the northern limit of breeding ranges and suggest that further formal survey effort will undoubtedly lead to additional new breeding records.

Isotopic spiking and food dye experiments provide evidence that nestling Canada Jays (Perisoreus canadensis) receive cached food from their parents

While many animals rely on stored food to survive periods of no or few resources, some of these species may also use cached food to feed young. The Canada Jay (Perisoreus canadensis (Linnaeus, 1766)) is a territorial, food-caching resident of North American boreal forests. Canada Jays have high winter survival when fresh food is rarely available and achieve high fledging rates before the return and breeding of most sympatric migratory passerines. Stored food must account for the Canada Jay’s winter survival, but it is less certain that stores are also used to feed nestlings. We conducted two experiments seeking evidence of such use, one using food marked with dye and a second using 15N-enriched glycine. Breeding pairs were supplemented with marked food in the pre-incubation period, followed by our attempts to find the marker in nestling feces (dye experiment) or feathers (15N-enrichment experiment). Both experiments provided evidence that at least some nestlings had ingested the marker, although the importance of this behaviour within the population remains unknown. Our study demonstrates how food markers can be used to infer feeding behaviour that has not been directly recorded in the wild, even after over 50 years of close observation.

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.

A test of the predator avoidance hypothesis to explain delayed onset of communal breeding

A neglected question in the study of communal breeding concerns why alloparental behaviour begins at variously late stages in the breeding cycle. In group-living corvids, the delay tends to be longer in species that are small and (or) typically have only a small nonbreeder complement. This pattern has been attributed to the relatively poor defensive capabilities of such species and their consequently greater need to minimize predator-attracting traffic to the nest or fledglings. We tested this predator avoidance hypothesis with the Canada Jay (Perisoreus canadensis (Linnaeus, 1766)), a species in which the feeding of young by any nonbreeders in the family group is delayed until the fledgling period. We reasoned that, on Anticosti Island, Quebec (Canada), in the absence of squirrels and other nest predators, nonbreeders might be permitted to feed nestlings as well as fledglings, and that breeders might feed nestlings more frequently (with smaller food loads) than on the mainland. We found no evidence for either prediction and thus no support for the predator avoidance hypothesis but suggest that Anticosti Canada Jays may have had insufficient time to evolve behaviour more appropriate for their predator-free environment. Secondarily, we confirmed that in all observed instances, the nonbreeders were offspring of the breeding pair from previous years and that they therefore failed to provision nestlings in spite of an apparent genetic interest to do so.

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.

Multilocus genetic analyses and spatial modeling reveal complex population structure and history in a widespread resident North American passerine (Perisoreus canadensis).

An increasing body of studies of widely distributed, high latitude species shows a variety of refugial locations and population genetic patterns. We examined the effects of glaciations and dispersal barriers on the population genetic patterns of a widely distributed, high latitude, resident corvid, the gray jay (Perisoreus canadensis), using the highly variable mitochondrial DNA (mtDNA) control region and microsatellite markers combined with species distribution modeling. We sequenced 914 bp of mtDNA control region for 375 individuals from 37 populations and screened seven loci for 402 individuals from 27 populations across the gray jay range. We used species distribution modeling and a range of phylogeographic analyses (haplotype diversity, ΦST, SAMOVA, F ST, Bayesian clustering analyses) to examine evolutionary history and population genetic structure. MtDNA and microsatellite markers revealed significant genetic differentiation among populations with high concordance between markers. Paleodistribution models supported at least five potential areas of suitable gray jay habitat during the last glacial maximum and revealed distributions similar to the gray jay's contemporary during the last interglacial. Colonization from and prolonged isolation in multiple refugia is evident. Historical climatic fluctuations, the presence of multiple dispersal barriers, and highly restricted gene flow appear to be responsible for strong genetic diversification and differentiation in gray jays.

Reduced reproductive performance associated with warmer ambient temperatures during incubation in a winter‐breeding, food‐storing passerine

Timing of reproduction can influence individual fitness whereby early breeders tend to have higher reproductive success than late breeders. However, the fitness consequences of timing of breeding may also be influenced by environmental conditions after the commencement of breeding. We tested whether ambient temperatures during the incubation and early nestling periods modulated the effect of laying date on brood size and dominant juvenile survival in gray jays (Perisoreus canadensis), a sedentary boreal species whose late winter nesting depends, in part, on caches of perishable food. Previous evidence has suggested that warmer temperatures degrade the quality of these food hoards, and we asked whether warmer ambient temperatures during the incubation and early nestling periods would be associated with smaller brood sizes and lower summer survival of dominant juveniles. We used 38 years of data from a range‐edge population of gray jays in Algonquin Provincial Park, Ontario, where the population has declined over 50% since the study began. Consistent with the “hoard‐rot” hypothesis, we found that cold temperatures during incubation were associated with larger brood sizes in later breeding attempts, but temperatures had little effect on brood size for females breeding early in the season. This is the first evidence that laying date and temperature during incubation interactively influence brood size in any bird species. We did not find evidence that ambient temperatures during the incubation period or early part of the nestling period influenced summer survival of dominant juveniles. Our findings provide evidence that warming temperatures are associated with some aspects of reduced reproductive performance in a species that is reliant on cold temperatures to store perishable food caches, some of which are later consumed during the reproductive period.

Male experience buffers female laying date plasticity in a winter-breeding, food-storing passerine

Phenotypic plasticity allows individuals to adjust reproductive timing in response to variation in the environment but little is known about how other factors, such as habitat quality, social environment and experience, may influence adjustments in the timing of breeding. We evaluated intrinsic (female age), environmental and social factors influencing laying date plasticity and assessed the effect of laying date on reproductive success in a population of grey jays, Perisoreus canadensis , over nearly four decades (1978–2015). Grey jays rely on stored food during their late-winter nesting season, a unique life history context to study plasticity in reproductive timing. Overall, females tended to lay eggs earlier in response to higher prelaying temperatures and advanced laying date at similar rates over their lives. Male age interacted with both temperature and female age to influence laying date. Females mated to older males were more likely to breed earlier at lower temperatures than females mated to younger males but there was little effect of male age under warmer conditions. Similarly, younger females mated to older males were more likely to breed earlier than younger females mated to younger males but there was little effect of male age when females were older. Across all years, earlier laying relative to other breeders in the population led to higher probability of nest success and summer survival for dominant juveniles. Our results suggest that individual females adjust laying date in response to temperature and provide the first evidence that male experience plays an important, and probably underappreciated, role in how females adjust their timing of breeding over their lives and with respect to annual variation in the environment.

Experimental evidence and 43 years of monitoring data show that food limits reproduction in a food-caching passerine.

Several species of birds and mammals overcome periods of scarcity by caching food, but for the vast majority of species, it is virtually unknown whether they are food limited during these periods. The Gray Jay (Perisoreus canadensis) is a boreal-resident, food-caching passerine that breeds in late winter when fresh food is scarce. Using a two-year experiment and 43 years of monitoring data, we examined the food limitation hypothesis in a population of Gray Jays in Algonquin Park, Ontario, Canada, that has declined by over 50% in the last three decades. Breeding pairs that were experimentally food supplemented during the pre-breeding period laid eggs earlier in the season and had larger brood sizes than non-supplemented controls. From the long-term data, we found strong evidence that pairs that were regularly supplemented by the public (park visitors) tended to lay eggs earlier and have larger clutches and brood sizes compared to pairs that were not supplemented. Nestling body condition (mass controlled for body size) was not influenced by either experimental or public food supplementation. Our results support the hypothesis that Gray Jays are food limited during their late-winter breeding period and suggest that warmer fall temperatures, which have been hypothesized to lead to cache spoilage, may have a significant impact on reproductive success in this declining population. Moreover, our results contribute to understanding how public feeding can influence the fitness of wild animals.

An example of phenotypic adherence to the island rule? - Anticosti gray jays are heavier but not structurally larger than mainland conspecifics.

The island rule refers to the tendency of small vertebrates to become larger when isolated on islands and the frequent dwarfing of large forms. It implies genetic control, and a necessary linkage, of size and body-mass differences between insular and mainland populations. To examine the island rule, we compared body size and mass of gray jays (Perisoreus canadensis) on Anticosti Island, Québec, located in the Gulf of St. Lawrence, with three mainland populations (2 in Québec and 1 in Ontario). Although gray jays on Anticosti Island were ca 10% heavier, they were not structurally larger, than the three mainland populations. This suggests that Anticosti jays are not necessarily genetically distinct from mainland gray jays and that they may have achieved their greater body masses solely through packing more mass onto mainland-sized body frames. As such, they may be the first-known example of a proposed, purely phenotypic initial step in the adherence to the island rule by an insular population. Greater jay body mass is probably advantageous in Anticosti's high-density, intensely competitive social environment that may have resulted from the island's lack of mammalian nest predators.

Linking the availability of cached food to climate change: an experimental test of the hoard-rot hypothesis

Understanding how long-term changes in climate influence populations requires knowledge of the mechanisms by which climate influences individual success and population abundance. We explored the implications of warmer falls and winters on a nonmigratory, food-caching bird, the Gray Jay (Perisoreus canadensis (L., 1766)), in Algonquin Park, Ontario, by experimentally examining the influence of temperature on the caloric content and mass of simulated cached food. Using three sites in, north of, and south of Algonquin, we examined the hypothesis that warmer temperatures would negatively influence the preservation of cached food by accelerating decay. Food cached at the highest latitude site retained the greatest mass and caloric content over 180 days of exposure, but there was no difference between the intermediate and the low latitude sites. We also show that Algonquin experienced a winter climate similar to our northernmost site as recently as 1990, and our results suggest that food availability in Algonquin has likely decreased since then, due to a warming climate. Interestingly, this coincides with a decline in territory occupancy by Gray Jays in Algonquin. Our results point to a unique mechanism by which climate warming may influence the persistence of a boreal species through its influence on winter food availability.

Analysis of combined data sets yields trend estimates for vulnerable spruce-fir birds in northern United States

Continental-scale monitoring programs with standardized survey protocols play an important role in conservation science by identifying species in decline and prioritizing conservation action. However, rare, inaccessible, or spatially fragmented communities may be underrepresented in continental-scale surveys. Data on these communities often come from decentralized, local monitoring efforts that differ in their goals and survey protocols. We combine 16 point count datasets, controlling for differences in protocol and detection probabilities to estimate regional trends for 14 spruce-fir forest bird species across Northeastern and Midwestern United States, a vulnerable community threatened by numerous anthropogenic stressors and widely considered a priority for conservation. Our analyses indicated that four species considered as ecological indicators for this community, Bicknell’s Thrush (Catharus bicknelli), Magnolia Warbler (Setophaga magnolia), Blackpoll Warbler (Setophaga striata) and Yellow-bellied Flycatcher (Empidonax flaviventris), each exhibited significant declines. Olive-sided Flycatcher (Contopus cooperi), a species of concern in parts of its range, and two additional species for which no previous concern existed, the Evening Grosbeak (Coccothruastes vespertinus) and the Gray Jay (Perisoreus canadensis), each also showed significant overall declines. Five out of nine species with sufficient data for analyses from Northeastern and Midwestern surveys showed significant differences in trends between these regions. Spruce-fir obligate species were more likely to decline significantly than species that use spruce-fir in addition to other habitat types. These results demonstrate the value of combining disparate data sources for analyzing regional patterns of population trends to confirm and extend conservation concern for some species and identify others for which additional attention may be needed.

Recreation changes the use of a wild landscape by corvids

ABSTRACT As urban areas have grown in population, use of nearby natural areas for outdoor recreation has also increased, potentially influencing bird distribution in landscapes managed for conservation. Members of the family Corvidae (crows, ravens, jays, and magpies) have strong interactions with humans and may be directly affected by recreation in wild landscapes. In Mount Rainier National Park, we evaluated the effects of vegetation, visitor use, and the availability of human-subsidized food on the use of landscape features by 4 corvid species: Steller's Jay (Cyanocitta stelleri), Gray Jay (Perisoreus canadensis), Common Raven (Corvus corax), and Clark's Nutcracker (Nucifraga columbiana). We conducted >1,400 point counts across areas that varied in habitat and in the degree of human recreational impact. We calculated predicted occupancy values while allowing for variation in detection probability. In addition to species-specific suites of vegetation and landscape variables, we found that patterns of hu...

Causes and consequences of pre‐laying weight gain in a food‐caching bird that breeds in late winter

Animals that reside at high latitudes and altitudes year‐round often use cached food to survive over the winter months, but a few species also rely on stored food to sustain them during the breeding season when the nutritional requirements of females are higher than normal. Gray jays Perisoreus canadensis rely on perishable cached food during the winter and females begin breeding in late winter when fresh food is rarely available. To examine pre‐laying patterns of weight gain, as well as the causes and consequences of weight gain among individuals, we weighed females regularly throughout the pre‐laying period. Females began increasing their weight approximately nine days prior to their first egg date, and on average increased their body weight by 25%, which is on par with other bird species that rely on non‐cached food. Final pre‐laying weight was positively influenced by the percent of conifers on territories, providing some support for previous results showing that coniferous trees are better able to preserve cached food. We also found that both final pre‐laying weight and the rate of weight gain were positively related to female age, supporting the hypothesis that female caching ability improves with age. With increasing final weight, females tended to lay larger clutches and hatched more nestlings, despite the fact that final weight was not influenced by weight at the beginning of the weighing period. Our results confirm that gray jays are able to reach breeding condition while relying primarily on food stored before winter, and suggest a novel mechanism by which habitat‐mediated carry‐over effects and female age may influence reproductive performance in a food‐caching animal.

Contrasting patterns of survival and dispersal in multiple habitats reveal an ecological trap in a food-caching bird

A comprehensive understanding of how natural and anthropogenic variation in habitat influences populations requires long-term information on how such variation affects survival and dispersal throughout the annual cycle. Gray jays Perisoreus canadensis are widespread boreal resident passerines that use cached food to survive over the winter and to begin breeding during the late winter. Using multistate capture-recapture analysis, we examined apparent survival and dispersal in relation to habitat quality in a gray jay population over 34 years (1977-2010). Prior evidence suggests that natural variation in habitat quality is driven by the proportion of conifers on territories because of their superior ability to preserve cached food. Although neither adults (>1 year) nor juveniles (<1 year) had higher survival rates on high-conifer territories, both age classes were less likely to leave high-conifer territories and, when they did move, were more likely to disperse to high-conifer territories. In contrast, survival rates were lower on territories that were adjacent to a major highway compared to territories that did not border the highway but there was no evidence for directional dispersal towards or away from highway territories. Our results support the notion that natural variation in habitat quality is driven by the proportion of coniferous trees on territories and provide the first evidence that high-mortality highway habitats can act as an equal-preference ecological trap for birds. Reproductive success, as shown in a previous study, but not survival, is sensitive to natural variation in habitat quality, suggesting that gray jays, despite living in harsh winter conditions, likely favor the allocation of limited resources towards self-maintenance over reproduction.

High latitudes and high genetic diversity: Phylogeography of a widespread boreal bird, the gray jay (Perisoreus canadensis)

We describe range-wide phylogeographic variation in gray jays (Perisoreus canadensis), a boreal Nearctic corvid that occurs today primarily in recently glaciated regions. Phylogenetic analysis of mitochondrial DNA (1041 base pairs ND2 gene; N=205, 50 localities) revealed four reciprocally monophyletic groups. One widespread clade occurs across the North American boreal zone, from Newfoundland to Alaska and southwest into Utah. Three other clades occur at lower latitudes in the montane West in Colorado, the northern Rocky Mountains, and the Pacific Northwest respectively. The geographic distribution of clades in gray jays corresponds with a general pattern that is emerging for boreal taxa, having one widespread northern clade and one or more geographically restricted southwestern clades. Population genetic analyses indicate that the larger boreal clade is genetically structured and harbors significantly more genetic diversity than those clades occurring at lower latitudes. Species distribution modeling (SDM) revealed multiple putative Pleistocene refugia including several occurring at higher latitudes. We suggest that multiple post-glacial colonization routes, some of which originate from these northern refugia, are responsible for the relatively high genetic diversity at high latitudes. Conversely, lower latitude clades show little variation, probably as a result of historical restriction to smaller geographical areas with smaller long-term population sizes. This ‘upside-down’ pattern of genetic diversity contrasts with the conventional view that populations of north-temperate species occupying previously glaciated habitats should possess lower levels of diversity than their southern counterparts.

Experimental evidence for a novel mechanism driving variation in habitat quality in a food-caching bird

Variation in habitat quality can have important consequences for fitness and population dynamics. For food-caching species, a critical determinant of habitat quality is normally the density of storable food, but it is also possible that quality is driven by the ability of habitats to preserve food items. The food-caching gray jay (Perisoreus canadensis) occupies year-round territories in the coniferous boreal and subalpine forests of North America, but does not use conifer seed crops as a source of food. Over the last 33 years, we found that the occupancy rate of territories in Algonquin Park (ON, Canada) has declined at a higher rate in territories with a lower proportion of conifers compared to those with a higher proportion. Individuals occupying territories with a low proportion of conifers were also less likely to successfully fledge young. Using chambers to simulate food caches, we conducted an experiment to examine the hypothesis that coniferous trees are better able to preserve the perishable food items stored in summer and fall than deciduous trees due to their antibacterial and antifungal properties. Over a 1-4 month exposure period, we found that mealworms, blueberries, and raisins all lost less weight when stored on spruce and pine trees compared to deciduous and other coniferous trees. Our results indicate a novel mechanism to explain how habitat quality may influence the fitness and population dynamics of food-caching animals, and has important implications for understanding range limits for boreal breeding animals.