Black-capped Chickadees Research Articles

THE INFLUENCE OF SAMPLING DESIGN ON SPECIES TREE INFERENCE: A NEW RELATIONSHIP FOR THE NEW WORLD CHICKADEES (AVES:POECILE)

In this study, we explore the long-standing issue of how many loci are needed to infer accurate phylogenetic relationships, and whether loci with particular attributes (e.g., parsimony informativeness, variability, gene tree resolution) outperform others. To do so, we use an empirical data set consisting of the seven species of chickadees (Aves: Paridae), an analytically tractable, recently diverged group, and well-studied ecologically but lacking a nuclear phylogeny. We estimate relationships using 40 nuclear loci and mitochondrial DNA using four coalescent-based species tree inference methods (BEST, *BEAST, STEM, STELLS). Collectively, our analyses contrast with previous studies and support a sister relationship between the Black-capped and Carolina Chickadee, two superficially similar species that hybridize along a long zone of contact. Gene flow is a potential source of conflict between nuclear and mitochondrial gene trees, yet we find a significant, albeit low, signal of gene flow. Our results suggest that relatively few loci with high information content may be sufficient for estimating an accurate species tree, but that substantially more loci are necessary for accurate parameter estimation. We provide an empirical reference point for researchers designing sampling protocols with the purpose of inferring phylogenies and population parameters of closely related taxa.

If a bird flies in the forest, does an insect hear it?

Birds are major predators of many eared insects including moths, butterflies, crickets and cicadas. We provide evidence supporting the hypothesis that insect ears can function as 'bird detectors'. First, we show that birds produce flight sounds while foraging. Eastern phoebes (Sayornis phoebe) and chickadees (Poecile atricapillus) generate broadband sounds composed of distinct repetitive elements (approx. 18 and 20 Hz, respectively) that correspond to cyclic wing beating. We estimate that insects can detect an approaching bird from distances of at least 2.5 m, based on insect hearing thresholds and sound level measurements of bird flight. Second, we show that insects with both high and low frequency hearing can hear bird flight sounds. Auditory nerve cells of noctuid moths (Trichoplusia ni) and nymphalid butterflies (Morpho peleides) responded in a bursting pattern to playbacks of an attacking bird. This is the first study to demonstrate that foraging birds generate flight sound cues that are detectable by eared insects. Whether insects exploit these sound cues, and alternatively, if birds have evolved sound-reducing foraging tactics to render them acoustically 'cryptic' to their prey, are tantalizing questions worthy of further investigation.

The Effect of Polychlorinated Biphenyls on the Song of Two Passerine Species

Polychlorinated biphenyls (PCBs) are synthetic chemical pollutants with demonstrated detrimental toxic and developmental effects on humans and wildlife. Laboratory studies suggest that PCBs influence behavior due to their effects on endocrine and neurological systems, yet little is known about the behavioral consequences of sublethal PCB exposure in the field. Additionally, specific PCB congener data (in contrast to total PCB load) is necessary to understand the possible effects of PCBs in living organisms since number and position of chlorine substitution in a PCB molecule dictates the toxicity and chemical fate of individual PCB congeners. We non-lethally investigated total PCB loads, congener specific PCB profiles, and songs of black-capped chickadees (Poecile atricapillus) and song sparrows (Melospiza melodia) along a historical PCB gradient at the Hudson River in New York State. Our results indicate that black-capped chickadees and song sparrows have higher total blood PCBs in regions with higher historic PCB contamination. The two bird species varied substantially in their congener-specific PCB profiles; within sites, song sparrows showed a significantly higher proportion of lower chlorinated PCBs, while black-capped chickadees had higher proportions of highly chlorinated PCBs. In areas of PCB pollution, the species-specific identity signal in black-capped chickadee song varied significantly, while variation in song sparrow trill performance was best predicted by the mono-ortho PCB load. Thus, PCBs may affect song production, an important component of communication in birds. In conclusion, we suggest that the ramifications of changes in song quality for bird populations may extend the toxic effects of environmental PCB pollution.

Habitat Quality Affects Early Physiology and Subsequent Neuromotor Development of Juvenile Black-Capped Chickadees

In songbirds, the ability to learn and render the species-specific song is influenced by the development of both the song nuclei in the brain and the syrinx (bird's vocal apparatus) early in the bird's life. In black-capped chickadees (Poecille atricapillus), habitat quality is known to affect song structure, with birds in high-quality habitat (mature forest) having a higher song consistency than birds in low-quality habitat (young forest). Although this difference is suspected to stem from differences in development, the developmental status of juvenile birds in either habitat remains unexplored. In this study, we used ptilochronology and feather corticosterone to compare the conditional state of juvenile chickadees in young and mature forest during two distinct periods of song learning - the sensory phase, which occurs prior to settlement, and the sensorimotor phase, which occurs post-settlement. A sample of juvenile males was captured and euthanized several weeks prior to their first breeding season to compare the development of song center nuclei and syrinx in both habitats. The corticosterone levels of natally-grown feathers were greater among birds that settled in mature than young forests - as these feathers were grown pre-settlement, they reflect differences in physiology during the sensory phase. This difference in conditional state is reflected by differences in syrinx and song center nuclei development later during the sensorimotor phase - birds in young forest have smaller syrinx, and moderately-larger RA, than birds in mature forest. Those differences could be responsible for the difference in consistency in song structure observed across habitats. The difference in physiological state across habitats, combined with potential compounding effect of differences in winter resources between habitats, could influence the difference in syrinx and neural development seen in juvenile males during the early spring, and influence the male's ability to learn and render their species-specific song.

Heterospecific discrimination of Poecile vocalizations by zebra finches (Taeniopygia guttata).

Previous perceptual research with black-capped and mountain chickadees has demonstrated that the D note of the namesake chick-a-dee call controlled species-based discrimination compared to other note types in this call. In the current experiment, we sought to determine whether discrimination performance of the chickadees was controlled by stimulus-specific properties or due to learning through experience. To accomplish this, we tested zebra finches, a songbird species that is distantly related to chickadees, and also unfamiliar with black-capped and mountain chickadee vocalizations, on the same species-based discrimination on which black-capped and mountain chickadees were previously trained. We found that zebra finches learned the discrimination in the fewest number of trials with the D note, compared to other note types (i.e., the A, B, and C notes). In addition, we compared the current results to earlier work and found that zebra finches learned the discrimination in fewer trials compared to black-capped chickadees, and, across all species, males learned the discrimination in fewer trials than females. We discuss the roles that acoustic complexity and learning play in classification of the three species of songbirds tested. More generally, these results point to the benefits derived from testing members of each sex in species that vary in their natural history, vocal output, and phylogenetic relatedness as a means to uncover the mechanisms underlying acoustic communication. (PsycINFO Database Record (c) 2013 APA, all rights reserved).

Tree stocking affects winter bird densities across a gradient of savanna, woodland, and forest in the Missouri Ozarks

Savanna and woodland were historically prevalent in the midwestern United States, and managers throughout the area are currently attempting to restore these communities. Better knowledge of the responses of breeding and non-breeding birds to savanna and woodland restoration is needed to inform management.We surveyed abundance of winter resident birds across a gradient of tree stocking encompassed by savannas, woodlands, and non-managed forests in the Missouri Ozark Highlands, USA, and assessed the effect of stocking on bird densities. We conducted point counts at 560 locations along 42 transects on 10 sites across 2 winters between December and February, 2009-2011. We estimated detection probabilities and densities of birds using hierarchical, distance-based abundance models that incorporate covariates into estimation of both detection probabilities and densities.We fit models for 12 species with >49 detections and met model assumptions. Detection probabilities were affected by observer, temperature, wind speed, and stocking, and there was some level of support of the effects of stocking on the densities of all 12 species. Densities of black-capped plus Carolina chickadees (Poecile atricapillus, P. carolinensis), brown creeper (Certhia americana), golden-crowned kinglet (Regulus satrapa), and red-bellied woodpecker (Melanerpes carolinus) increased with stocking; whereas, densities of eastern bluebird (Sialia sialis), hairy woodpecker (Picoides villosus), and northern flicker (Colaptes auratus) were greatest at intermediate values of stocking. Although densities of 7 of the 12 species varied substantially across the range of stocking, all species used all 3 communities across a wide range of stocking. We provide winter densities of 12 of the most common overwintering bird species in the Missouri Ozarks; the relationships between their densities and stocking can be used to guidemanagement where managers have bird-species-specific objectives.

Habitat-induced changes in song consistency affect perception of social status in male chickadees

The ability of male black-capped chickadees to maintain consistent internal structure between successive iterations of their songs is affected by both their social rank and the quality of their habitat. Lab studies reveal that female chickadees discriminate between songs of dominant and subordinate males, which vary in acoustic structure. We investigate whether males also rely on acoustic structure to assess rival quality during agonistic interactions, and whether habitat-induced differences in song consistency affect the perception of male rank. We conducted a playback experiment to simulate territorial intrusions by dominant, stimuli males into the territories of dominant, subject males; stimuli males were recorded either in low-quality (young forest) or high-quality (mature forest) habitats. Stimuli from low-quality habitat had lower song consistency than those from high-quality habitats, despite being recorded from males of equivalent social rank. Subject males for playbacks (also socially dominant males) were chosen from either habitat type. Subject males in mature forest responded less to young-forest stimuli compared to mature-forest stimuli, despite the stimuli in both cases being recorded from dominant males. Conversely, male subjects in young forest did not differentiate between stimuli, but their response to both stimuli was lower than that of mature-forest subject males to mature-forest stimuli. We demonstrate that the ability to maintain internal song structure in the black-capped chickadee constitutes a signal that appears to be used by males to assess the level of threat of intruders, and that this perception is affected by habitat from which the stimulus males were recorded.

Intra-Seasonal Flexibility in Avian Metabolic Performance Highlights the Uncoupling of Basal Metabolic Rate and Thermogenic Capacity

Stochastic winter weather events are predicted to increase in occurrence and amplitude at northern latitudes and organisms are expected to cope through phenotypic flexibility. Small avian species wintering in these environments show acclimatization where basal metabolic rate (BMR) and maximal thermogenic capacity (MSUM) are typically elevated. However, little is known on intra-seasonal variation in metabolic performance and on how population trends truly reflect individual flexibility. Here we report intra-seasonal variation in metabolic parameters measured at the population and individual levels in black-capped chickadees ( Poecile atricapillus ). Results confirmed that population patterns indeed reflect flexibility at the individual level. They showed the expected increase in BMR (6%) and MSUM (34%) in winter relative to summer but also, and most importantly, that these parameters changed differently through time. BMR began its seasonal increase in November, while MSUM had already achieved more than 20% of its inter-seasonal increase by October, and declined to its starting level by March, while MSUM remained high. Although both parameters co-vary on a yearly scale, this mismatch in the timing of variation in winter BMR and MSUM likely reflects different constraints acting on different physiological components and therefore suggests a lack of functional link between these parameters.

Reproductive isolation and cryptic introgression in a sky island enclave of Appalachian birds

AbstractReproductive isolation is central to the speciation process, and cases where the strength of reproductive isolation varies geographically can inform our understanding of speciation mechanisms. Although generally treated as separate species, Black‐capped chickadees (Poecile atricapillus) and Carolina chickadees (P. carolinensis) hybridize and undergo genetic introgression in many areas where they come into contact across the eastern United States and in the northern Appalachian Mountains. The Great Smoky Mountains harbor the last large breeding population of atricapillus in the southern Appalachians, isolated from the species’ main range by nearly 200 km. This population is believed to be reproductively isolated from local carolinensis due to an unusual, behaviorally mediated elevational range gap, which forms during the breeding season and may function as an incipient reproductive isolating mechanism. We examined the effectiveness of this putative isolating mechanism by looking for genetic introgression from carolinensis in Great Smoky Mountain atricapillus. We characterized this population and parental controls genetically using hundreds of amplified fragment length polymorphism (AFLP) loci as well as mitochondrial DNA (mtDNA) sequence data from cytochrome‐b. Great Smoky Mountain atricapillus have experienced nuclear genetic introgression from carolinensis, but at much lower levels than other populations near the hybrid zone to the north. No mitochondrial introgression was detected, in contrast to northern contact areas. Thus, the seasonal elevational range gap appears to have been effective in reducing gene flow between these closely related taxa.

Avian Incubation Patterns Reflect Temporal Changes in Developing Clutches

Incubation conditions for eggs influence offspring quality and reproductive success. One way in which parents regulate brooding conditions is by balancing the thermal requirements of embryos with time spent away from the nest for self-maintenance. Age related changes in embryo thermal tolerance would thus be expected to shape parental incubation behavior. We use data from unmanipulated Black-capped Chickadee (Poecile atricapillus) nests to examine the temporal dynamics of incubation, testing the prediction that increased heat flux from eggs as embryos age influences female incubation behavior and/or physiology to minimize temperature fluctuations. We found that the rate of heat loss from eggs increased with embryo age. Females responded to increased egg cooling rates by altering incubation rhythms (more frequent, shorter on- and off- bouts), but not brood patch temperature. Consequently, as embryos aged, females were able to increase mean egg temperature and decrease variation in temperature. Our findings highlight the need to view full incubation as more than a static rhythm; rather, it is a temporally dynamic and finely adjustable parental behavior. Furthermore, from a methodological perspective, intra- and inter-specific comparisons of incubation rhythms and average egg temperatures should control for the stage of incubation.

Character displacement in dawn chorusing behaviour of sympatric mountain and black-capped chickadees

Signals of closely related species tend to be more distinct when occurring in sympatry than in allopatry. Such differences allow species-specific identification and presumably reduce interspecific mating. Among chickadees, dawn chorus behaviour affects female mate choice. Within our sympatric study population of black-capped chickadees, Poecile atricapillus, and mountain chickadees, Poecile gambeli, directional hybridization occurs via extrapair matings between female mountain chickadees (the subordinate species) and male black-capped chickadees (the dominant species). In this study, we compared dawn singing from this sympatric population with dawn recordings from allopatric populations of each species. Mountain chickadees used more chick-a-dee calls than songs during the dawn chorus when they co-occurred with black-capped chickadees in the sympatric population, but used similar proportions of calls and songs in the allopatric populations. We also found differences in the fine structure of the song; both species typically had a descending first note in their song (glissando), but mountain chickadees in the sympatric population used an ascending first note. The internote ratio between the first two notes of the song of the sympatric mountain chickadees lacked a characteristic frequency drop found in the allopatric mountain chickadee population and in the allopatric and sympatric populations of black-capped chickadees. Geographical analysis of songs of mountain chickadees across western North America revealed consistent differences in song features among sympatric/allopatric populations in different regions, but the nature of character shifts were not always parallel among populations. These findings illustrate possible character displacement in a subordinate species (mountain chickadee) to reduce acoustic overlap with a dominant heterospecific (black-capped chickadee).

Female song in black-capped chickadees (Poecile atricapillus): Acoustic song features that contain individual identity information and sex differences

In temperate songbirds, song has traditionally been considered a vocalization mainly produced by males. However, in many temperate species, it is now recognized that both males and females produce song. The function and structure of male black-capped chickadee (Poecile atricapillus) fee-bee song have been well-studied, but female song, although briefly described in some field reports, has not often been quantitatively examined. We recorded spontaneous fee-bee songs produced by wild-caught male and female adult black-capped chickadees housed in captivity and used bioacoustic analyses to examine seven acoustic features in the songs. Using potential for individual coding and discriminant function analyses, we found that songs were individually distinctive. Using additional discriminant function analyses, we found that songs could be correctly classified based on sex of the producer. Specifically, our results indicate that the frequency decrease within the fee note (i.e., fee glissando) varies between the songs produced by males and females and this suggests that the fee glissando may be used as a sex identifier. While this study provides a quantitative description of the acoustic structure of female song, the development, perception and function of female song in this species requires further examination.

Variation in hippocampal glial cell numbers in food‐caching birds from different climates

Enhancements to memory are associated with enhanced neural structures that support those capabilities. A great deal of work has examined this relationship in the context of natural variation in spatial memory capability and hippocampal (Hp) structure. Most studies have focused on volumetric and neuron measures, but have seldom examined the role of glial cells. Once considered involved only in supportive functions associated with neurons, the importance of glial cells in cognitive processes, including memory, is gaining more attention. Building upon our previous study on the relationship between the brain, memory, and environmental severity in food-caching birds, we compared the total number of Hp glial cells in wild-sampled and in lab-reared (common garden) black-capped chickadees (Poecile atricapillus) originating from two different environmental extremes. We found that birds from more harsh climate tended to have significantly more Hp glial cells than those from more mild climate and that lab-reared chickadees had significantly fewer Hp glial cells compared to the wild-sampled birds. These results suggest that population differences in glial numbers may be controlled, at least in part, by heritable mechanisms, but glial numbers appear to be additionally regulated by an individual's environment. The pattern of Hp glial cell abundance among our treatment groups closely followed that of the Hp volume, suggesting that Hp glial cell number may be associated with the Hp volume. Unlike Hp neurons, however, the number of Hp glial cells may be, at least in part, affected by an individual's experiences and environment.

Vector-Host Interactions and Epizootiology of Eastern Equine Encephalitis Virus in Massachusetts

Eastern equine encephalitis (EEE) virus is a highly pathogenic mosquito-borne zoonosis that is responsible for outbreaks of severe disease in humans and equines, resulting in high mortality or severe neurological impairment in most survivors. In the northeastern United States, EEE virus is maintained in an enzootic cycle involving the ornithophilic mosquito, Culiseta melanura (Coquillett) and passerine birds in freshwater swamp habitats. To evaluate the role of Cs. melanura and Culiseta morsitans (Theobald) in recent episodes of EEE virus activity in Massachusetts, we collected blood-fed mosquitoes between June, 2007, and October, 2008, from virus foci in 6 counties, and identified the source of blood meals by PCR amplification of mitochondrial cytochrome b gene and sequencing. Analysis of 529 Cs. melanura and 25 Cs. morsitans revealed that nearly 99% and 96% of mosquitoes, respectively, acquired blood meals solely from avian hosts. American Robin, Turdus migratorius Linnaeus was identified as the most common vertebrate host for Cs. melanura (21.7%, n=115), followed by Tufted Titmouse, Baeolophus bicolor (L.) (8.7%, n=46), Black-capped Chickadee, Poecile atricapillus (L.) (8.5%, n=45), Scarlet Tanager, Piranga olivacea (Gmelin) (6.8%, n=36), Field Sparrow, Spizella pusilla (Wilson) (6.2%, n=33), Northern Cardinal, Cardinalis cardinalis (L.) (5.7%, n=30), and other mostly Passeriformes birds. Mammalian-derived blood meals were identified as white-tailed deer, Odocoileus virginianus Zimmermann, domestic cow, Bos taurus L., and human, Homo sapiens L. There were 4 isolations of EEE virus, West Nile virus, and Highland J virus from Cs. melanura. Our results in conjunction with other lines of evidence, including reservoir competency, prevalence of antibody, and infection in nature, suggest that the American Robin, Tufted Titmouse, Black-capped Chickadee, and a few other passerine birds may play key roles in supporting EEE virus transmission in Massachusetts. Infrequent blood feeding of Cs. melanura on mammalian hosts, including humans, also indicates that this mosquito may occasionally contribute to epidemic/epizootic transmission of EEE virus in this region.

Macroscopic, Histologic, and Ultrastructural Lesions Associated With Avian Keratin Disorder in Black-Capped Chickadees (Poecile atricapillus)

An epizootic of beak abnormalities (avian keratin disorder) was recently detected among wild birds in Alaska. Here we describe the gross, histologic, and ultrastructural features of the disease in 30 affected adult black-capped chickadees (Poecile atricapillus). Grossly, there was elongation of the rhamphotheca, with varying degrees of lateral deviation, crossing, and gapping between the upper and lower beak. Not uncommonly, the claws were overgrown, and there was alopecia, scaling, and crusting of the skin. The most prominent histopathologic features in the beak included epidermal hyperplasia, hyperkeratosis, and core-like intrusions of necrotic debris. In affected birds, particularly those with moderate to severe beak overgrowth, there was remodeling of premaxillary and mandibular bones and various dermal lesions. Lesions analogous to those found in beaks were present in affected claws, indicating that this disorder may target both of these similar tissues. Mild to moderate hyperkeratosis occurred in other keratinized tissues, including skin, feather follicles, and, occasionally, sinus epithelium, but typically only in the presence of microbes. We did not find consistent evidence of a bacterial, fungal, or viral etiology for the beak lesions. The changes observed in affected birds did not correspond with any known avian diseases, suggesting a potentially novel hyperkeratotic disorder in wild birds.

Chickadees fail standardized operant tests for octave equivalence

Octave equivalence occurs when an observer judges notes separated by a doubling in frequency perceptually similar. The octave appears to form the basis of pitch change in all human cultures and thus may be of biological origin. Previously, we developed a nonverbal operant conditioning test of octave generalization and transfer in humans. The results of this testing showed that humans with and without musical training perceive the octave relationship between pitches. Our goal in the current study was to determine whether black-capped chickadees, a North American songbird, perceive octave equivalence. We chose these chickadees because of their reliance on pitch in assessing conspecific vocalizations, our strong background knowledge on their pitch height perception (log-linear perception of frequency), and the phylogenetic disparity between them and humans. Compared to humans, songbirds are highly skilled at using pitch height perception to classify pitches into ranges, independent of the octave. Our results suggest that chickadees used that skill, rather than octave equivalence, to transfer the note-range discrimination from one octave to the next. In contrast, there is evidence that at least some mammals, including humans, do perceive octave equivalence.

Shift of song frequencies in response to masking tones

Ambient noise can interfere with signal transmission and detection across many taxa and modalities. Evidence suggests that, over time, signals evolve to minimize interference from ambient noise and other signalling animals. Less well studied is the possibility of short-term behavioural responses to transient ambient noise, in which animals actively adjust signal parameters to recover signalling efficacy. Here we test animals' capacity to adjust vocal signal parameters in the face of transient acoustic interference. In field trials we monitored the songs of territorial male black-capped chickadees, Poecile atricapillus, determined the frequencies of their ‘fee-bee’ songs, and broadcast tones to closely mask subjects' ‘bee’ notes. We also presented control nonmasking tones of 5 kHz, well above birds' song frequencies. Our main finding was that males responded to masking tones by shifting song frequencies after an average of 66.4 s from tone onset, whereas frequency shifts in the presence of nonmasking tones occurred only after an average of 95.8 s. The quicker shift in frequencies in the face of masking noise provides new evidence for vocal behavioural plasticity, and further reveals how behavioural plasticity together with evolutionary adaptations can minimize the detrimental effects of ambient noise on communication.

Dominance and geographic information contained within black-capped chickadee (Poecile atricapillus) song

In songbirds, male song is an acoustic signal used to attract mates and defend territories. Typically, song is an acoustically complex signal; however, the fee-bee song of the black-capped chickadee is relatively simple. Despite this relative simplicity, two previous studies (Christie et al., 2004b; Hoeschele et al., 2010) found acoustic features within the fee-bee song that contain information regarding an individual’s dominance rank; however each of these studies reported a different dominance-related acoustic cue. Specifically, the relative amplitude of the two notes differed between the songs of dominant and subordinate males from northern British Columbia, while the interval pitch ratio differed between the songs of dominant and subordinate males from eastern Ontario. In the current study, we examined six acoustic features within songs from both of the chickadee populations (northern British Columbia and eastern Ontario) examined in these previous studies and used bioacoustic analyses and discriminant function analyses to determine whether there is a consistent dominance-related acoustic cue across both, or in each of these populations. Consistent with the previous findings, the current results indicate that relative amplitude differs based on dominance status in the songs from British Columbia; however, our results failed to reach significance with songs from Ontario. These results suggest that acoustic cues that signal a male’s dominance in this species vary with geographic location. Furthermore, examining songs from these two locations and one additional location in northern British Columbia, we found that discriminant function analyses could correctly classify songs based on geographic location. Considering the broad extent of the species’ range, black-capped chickadee song is considered relatively invariant; however, our results suggest that there is geographic variation in songs, although the differences are subtle compared to geographic song variation in other species.

Phenotype manipulations confirm the role of pectoral muscles and haematocrit in avian maximal thermogenic capacity

In small resident bird species living at northern latitudes, winter cold acclimatization is associated with an increase in pectoral muscle size and haematocrit level, and this is thought to drive the seasonal increase in summit metabolic rate (Msum, a measure of maximal shivering thermogenic capacity). However, evidence suggesting that pectoral muscle size influences Msum is correlational and the link between haematrocrit level and Msum remains to be demonstrated. We experimentally tested the relationship between pectoral muscle size and Msum by manipulating muscle size using a feather clipping protocol in free-living wintering black-capped chickadees (Poecile atricapillus). This also allowed us to investigate the link between haematocrit and thermogenic capacity. After a first series of measures on all birds, we cut half of the flight feathers of experimental individuals (N=14) and compared their fat and pectoral muscle scores, Msum and haematocrit level at recapture with their previous measures and with those of control birds (N=17) that were captured and recaptured at comparable times. Results showed that: (1) experimental birds developed larger pectoral muscles than control individuals and (2) mass-independent Msum was up to 16% higher in birds expressing large pectoral muscles. Msum was also positively correlated with haematocrit, which was not affected by the experimental manipulation. These findings demonstrate that, for a given body mass, large pectoral muscles are associated with a higher Msum in black-capped chickadees and that oxygen carrying capacity likely supports thermogenesis in this species.

Differential hippocampal gene expression is associated with climate‐related natural variation in memory and the hippocampus in food‐caching chickadees

There is significant and often heritable variation in cognition and its underlying neural mechanisms, yet specific genetic contributions to such variation are not well characterized. Black-capped chickadees present a good model to investigate the genetic basis of cognition because they exhibit tremendous climate-related variation in memory, hippocampal morphology and neurogenesis rates throughout the North American continent, and these cognitive traits appear to have a heritable basis. We examined the hippocampal transcriptome profiles of laboratory-reared chickadees from the two most divergent populations to test whether differential gene expression in the hippocampus is associated with population differences in spatial memory, hippocampal morphology and adult hippocampal neurogenesis rates. Using high-resolution mRNA sequencing coupled to a de novo transcriptome assembly, we generated 23 295 consensus sequences, which predicted 16 206 protein sequences with 13 982 showing high similarity to known protein sequences or conserved hypothetical proteins in other species. Of these, we identified differential expression in nearly 380 genes, with 47 genes specifically linked to neurogenesis, apoptosis, synaptic function, and learning and memory processes. Many of the other differentially expressed genes, however, may be associated with other functions. Our study presents the first avian hippocampal transcriptome, and it is the first study identifying differential gene expression associated with natural variation in cognition and the hippocampus. Our results provide additional support to the hypothesis that population differences in memory, hippocampal morphology and neurogenesis in chickadees have likely resulted from natural selection that appears to act on memory and its underlying neural mechanisms.