Overlapping and matching in the song contests of black-capped chickadees

During vocal contests, animals alter both the timing and the patterning of responses to their opponents. Time-specific responses (such as overlapping an opponent's song) and pattern-specific responses (such as matching the type of an opponent's song) may reveal information about interacting animals. Here we explore the consequences of overlapping and frequency matching during song contests of male black-capped chickadees, Poecile atricapillus. Using interactive playback, we engaged birds of high and low dominance status in vocal interactions with a simulated territorial intruder. The playback intruder either overlapped or avoided overlapping the subjects' songs and either matched or avoided matching the frequency of the subjects' songs. Individuals who were overlapped by the playback intruder showed higher variability in their song length and song timing than individuals who were not overlapped. Individuals who were frequency-matched by the playback intruder responded with more agitated responses (more flights and passes over the speaker and closer distances of closest approach) and spent more time farther away from the loudspeaker. We argue that the timing of song delivery and the choice of song type are distinct functional components of vocal interactions, where overlapping and matching songs are threatening signals that have separate consequences for opponent behaviour and song performance. High-ranking males responded at greater distances from the loudspeaker in all treatments and responded with lower agitation levels than low-ranking males. We demonstrate that males of different quality show different behavioural responses to territorial intruders, where males of high status appear reluctant to engage an intruder as intensely as males of low status.     

Food hoarding behaviour of black-capped chickadees (Poecile atricapillus ) in relation to forest edges

In temperate forests, small birds avoid the use of forest edges in adverse winter weather suggesting high foraging costs in terms of energetic requirements. Since hoarding species will often retrieve caches during adverse winter weather, they may perceive forest edges, especially exposed ones, as low quality hoarding sites. We tested whether black-capped chickadees ( Poecile atricapillus) inhabiting fragmented forests modify and reduce hoarding activity near forest edges. We also tested whether hoarding behaviour will be most affected in sites with forest edges more exposed to extreme weather. Black-capped chickadees taking food from a feeder 30 m from the nearest forest edges hoarded items mostly towards the forest interior, whereas no preference in hoarding location was observed with birds taking food from a feeder placed >100 m from the edge. Furthermore, birds avoided direct flights towards forest edges and, at sites exposed to prevailing winds, hoarding trips were shorter than at other locations. These results suggest that individuals avoid hoarding near forest edges and there, they lower their investment in terms of hoarding effort. The observed difference in hoarding behaviour was more evident near forest edges delimiting wide unforested areas than in edges delimiting narrower unforested areas. Edge exposure to prevailing winds influenced hoarding behaviour much less. We suggest that hoarding birds may partially overcome the ecological costs of habitat loss and fragmentation due to abiotic edge effects. By hoarding food away from forest edges in good weather, they may use forest interiors as low-cost retrieval sites under adverse weather.     

Pitch-related cues in the songs of sympatric mountain and black-capped chickadees

Acoustic frequency (pitch) cues are known to be important in the recognition of conspecific song in a number of songbird species. Mountain chickadees (Poecile gambeli) and black-capped chickadees (Poecile atricapillus) are sympatric over parts of their ranges and their species-typical songs share many features. I examined the acoustic characteristics of song of these two congeners in a region of sympatry in southern Alberta, Canada. As reported for other populations in allopatry, black-capped chickadees emphasized relative frequency cues in song production. In particular, variation in the ratios between note frequencies was significantly less than variation in the note frequencies themselves. In contrast, songs of mountain chickadees did not have constant frequency ratios and contained an introductory acoustic element absent in black-capped chickadee song. Both species may rely on song note frequency or the presence of this introductory acoustic element when differentiating between conspecific song and heterospecific song. Song measures for chickadees in sympatry were similar to measures in allopatry, providing little evidence for character displacement in song production.     

Memory for food caches: not just for retrieval

Many animals use hoarding as a long-term strategy to ensurea food supply at times of shortage. Hoarders employ strategiesthat enhance their ability to relocate caches such as rememberingwhere caches are located. Long-term scatterhoarders, whose cacheshave potentially high pilferage rates, should also hoard ina way to reduce potential cache pilferers' ability to find caches.Previous studies have demonstrated that this could be achievedby hyperdispersing caches to reduce the foraging efficiencyof pilferers. This study investigates whether coal tits (Parusater) indeed place their caches away from existing ones. Inour experiment, birds hoarded food in 3 conditions: when cachesfrom a previous storage session were still present, when cachesfrom a previous storage session were not present anymore becausethe bird had retrieved them, and when caches from a previousstorage session had been removed by the experimenter. We showthat coal tits hoard away from existing caches and that theydo not use cues from extant caches to do this. This evidenceis consistent with the use of memory for the locations of previouscaches when deciding where to place new caches. This findinghas important implications for our understanding of the selectivepressures that have shaped spatial memory in food-hoarding birds.     

Honest advertisement and song output during the dawn chorus of black-capped chickadees

Costly signals can evolve under sexual selection, as only thosesignals that are difficult to produce and reflect the relativequality of individuals should be important in mate choice. Onesuch signal may be dawn singing behavior in birds. We assessedwhether the song output at dawn of breeding male black-cappedchickadees Parus atricapilhis honestly reflects quality, whererelative quality is assessed by relative dominance rank in winterflocks. Dawn choruses were recorded from 20 male chickadeesfrom 10 flocks during the fertile period of their mates in 1992,1994, and 1995. Dominance ranks of males were assessed by tabulatinginteractions at winter feeders from 1993 to 1995. A comparisonof the dawn singing behavior of the high-ranking and the low-rankingmales from each of the 10 flocks showed that high-ranking malesbegan singing earlier, sang longer, and sang at higher averageand maximum rates than low-ranking flockmates. Age of the maleshad less effect on song output at dawn than rank; older malestended to sing longer dawn choruses, but there was no differencein onset of singing, average song rate, or maximum song rateat dawn between hatch year and after-hatch year males. Our findingssuggest the dawn chorus can provide an accurate signal to femalesof the relative quality of their mate compared to neighboringmales     

Photoperiodic regulation of food storing and hippocampus volume in black-capped chickadees, Poecile atricapillus

In seasonal environments animals organize their behaviour around annual cycles of resource availability. Wild black-capped chickadees are most likely to hoard food in autumn. At this time of year chickadees are also reported to have a larger hippocampus, a brain area important for spatial memory. This study examined how photoperiodic condition affects these seasonal changes. Captive chickadees were exposed to one of three treatments. Photorefractory birds were held on long days (19:5 h light:dark) and had small gonads. Photosensitive birds were held on short days (LD 9:15 h) and also had small gonads. Photostimulated birds were switched from short to long days and quickly entered breeding condition with large gonads. Photosensitive birds (on short days) stored more seeds than photorefractory birds (on long days). Photostimulated birds stored seeds at a high rate when on short days, but reduced storing when transferred to long days. These results indicate that long days inhibit storing regardless of gonadal condition. There were no differences between groups in hippocampal volume, indicating that photoperiod can produce changes in food-storing behaviour without affecting hippocampal size. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.     

Seasonal variability in habitat structure may have shaped acoustic signals and repertoires in the black-capped and boreal chickadees

Many songbird species have evolved multiple vocalizations, or repertoires, that function to communicate various biological signals. More diverse repertoires may have evolved in response to the effects of seasonal variation in habitat structure on signal transmission. Such changes in habitat necessarily occur for migrating species, but they also occur for resident species that occupy deciduous forests. The black-capped chickadee (Poecile atricapillus) possesses a chick-a-dee call and a fee-bee song, but the closely related boreal chickadee (P. borealis) lacks a song. Consistent with the habitat variability hypothesis, the black-capped chickadee possesses a larger repertoire and primarily occupies deciduous forests, whereas the songless boreal chickadee occurs more often in coniferous forests. We explored the ecological basis of this hypothesis by recording audio playbacks of two species in two habitat types during two seasons. Specifically, we played both songs and calls of the black-capped chickadee and calls of the boreal chickadee in deciduous and coniferous habitats, prior to and after leaf-out. We measured attenuation and degradation in re-recorded vocalizations. For black-capped chickadees, the song was less degraded than the call in post-leaf, deciduous forests. The boreal chickadee call attenuated more quickly in all treatments, but maintained its acoustic structure better than both black-capped chickadee vocalizations in coniferous forests. Our results support the hypothesis that variable habitats provided a seasonal transmission benefit for both song and call in the black-capped chickadee, but that the transmission benefit of song is lost in the less variant coniferous forests, which may underlie the absence of a song in the boreal chickadee.     

Patterns of extrapair mating in relation to male dominance status and female nest placement in black-capped chickadees

In sexually promiscuous animals, females may benefit by nestingclose to the edge of their partner's territory to facilitateextrapair copulations. In the present study, we describe theextrapair mating system of black-capped chickadees, Poecileatricapillus, and test whether nest locations are influencedby conspecific attraction to extrapair partners. We conducteda spatial analysis of female mating strategies by using microsatellitepaternity analysis in conjunction with geographic informationsystem (GIS) analysis of nest and territory locations. Extrapairoffspring comprised 52 of 351 offspring (14.8%) and were presentin 19 of 57 broods (33.3%). Females paired to males with lowdominance status in the previous winter's flock hierarchy weremore likely to engage in a mixed reproductive strategy thanwere females paired to males with high dominance status. Femaleshad extrapair copulations and extrapair fertilizations withhigh-ranking males more often than with low-ranking males. Notall extrapair copulations resulted in extrapair fertilizations.Females constructed their nests within 16.8 ± 1.0 m ofthe edge of their partner's territory, significantly closerto the edge of their nearest neighbor's territory than to thecenter of their own partner's territory. Extrapair males usuallyshared territory boundaries with cuckolded males. Females pairedto low-ranking males constructed nests near the territory edgesof neighboring high-ranking males. However, females did nothave extrapair copulations with the neighbor nearest to theirnest or even with the high-ranking neighbor nearest to theirnest. We conclude that conspecific attraction to neighbors mayinfluence nesting location in black-capped chickadees; however,it does not operate by facilitating extrapair copulations.     

Dominant black-capped chickadees pay no maintenance energy costs for their wintering status and are not better at enduring cold than subordinate individuals

Winter requires physiological adjustments in northern resident passerines. Cold acclimatization is generally associated with an increase in physiological maintenance costs, measured as basal metabolic rate (BMR), and cold endurance, reflected by summit metabolic rate (M _sum). However, several northern species also form social groups in winter and a bird’s hierarchical position may influence the size of its metabolically active organs as well as its BMR. Winter metabolic performance in these species may therefore reflect a complex set of adjustments to both seasonal climatic variations and social environment. We studied the effect of social status on parameters of cold acclimatization (body mass, size of fat reserves and pectoral muscles, BMR and M _sum) in free-living black-capped chickadees (Poecile atricapillus). Birds that were structurally large and heavy for their body size, mostly dominant individuals, carried more fat reserves and had larger pectoral muscles. However, social status had little effect on metabolic performance in the cold. Indeed, M _sum was independent of social rank while mass-corrected BMR was slightly lower in dominant individuals, likely due to a statistical dilution effect caused by large metabolically inactive fat reserves. BMR and M _sum, whether considered in terms of whole-animal values, corrected for body mass or body size were nevertheless correlated, suggesting a functional link between these metabolic components. Our results therefore indicate that the energy cost of social dominance is not a generalized phenomenon in small wintering birds.     

Memory for the location of stored food in marsh tits

Marsh tits store several hundred food items per day, in separate locations within their territory, and recover most items within 24 h. Experiments in the laboratory determined that marsh tits return accurately to the sites of stored sunflower seeds 3 h and 24 h after storage. The proportion of returns exceeded chance encounter and initial preferences for those sites. The lack of interocular transfer in birds for some visual tasks provided a test for the hypothesis that memory is used to relocate stored food. After storing food with one eye covered, marsh tits did not return to storage sites when using only the ‘naive’ eye, but did so when using only the eye uncovered during storage. It seems likely that recovery of stored food by marsh tits and other birds occurs by memory of storage sites, not by chance encounter during foraging.     

Categorization and discrimination of "chick-a-dee" calls by wild-caught and hand-reared chickadees

Bloomfield and Sturdy [Bloomfield, L.L., Sturdy, C.B. All chick-a-dee calls are not created equally. Part I. Open-ended categorization by sympatric and allopatric chickadees. Behav. Proc., in press] previously reported that black-capped chickadees (Poecile atricapillus) discriminate conspecific from heterospecific (mountain chickadee, P. gambeli) 'chick-a-dee' calls, and their ability to accurately discriminate and classify the calls as belonging to separate species' defined categories was largely unaffected by their prior experience with mountain chickadees and their calls. To further examine the potential influence of experience on discrimination and categorization, we compare wild-caught black-capped chickadees, wild-caught mountain chickadees, and black-capped chickadees hand-reared among either adult laboratory-housed black-capped chickadees or adult laboratory-housed mountain chickadees on a true category/pseudo category chick-a-dee call discrimination task. Irrespective of group assignment, hand-reared birds performed as well as wild-caught birds and did not show a conspecific- or rearing-specific advantage in discrimination, categorization or memorization of chick-a-dee calls. While vocal learning is under the influence of ontogenetic experience, the results derived from the current methods suggest that experience (or a lack thereof) does not affect categorization and memorization abilities.     

Thiol and disulphide contents of hen ovalbumin. C-Terminal sequence and location of disulphide bond

1. The thiol and disulphide contents of hen ovalbumin were investigated by p-chloromercuribenzoate titration, by determination of cysteic acid content after performic acid oxidation, by measurement of uptake of radioactive iodoacetic acid, and by assay of S-aminoethylcysteine after reaction with ethyleneimine. All results showed that ovalbumin had 6 half-cystine residues. Experiments with and without reducing agents demonstrated that there were 4 thiol groups and 1 disulphide bond. 2. A peptide containing equimolar amounts of S-carboxymethyl-cysteine, serine, valine and proline, but no lysine or arginine, was obtained by radioactive labelling of the cysteine residues with iodo[¹⁴C]acetic acid followed by electrophoretic and chromatographic separation of tryptic digests. It was concluded that the C-terminal sequence of ovalbumin is -Cys-Val-Ser-Pro. 3. The location of the disulphide bond was studied by using a double-labelling technique. It was shown that one end of the disulphide was located in this C-terminal peptide.     

Maximizing feeding efficiency and minimizing time exposed to predators: a trade-off in the black-capped chickadee

Summary Animals often must feed away from protective cover, sometimes at a considerable risk of being preyed upon. Feeding at the maximum rate while away from cover may simultaneously minimize the time spent exposed to predators, but this is not always the case. Under some circumstances, carrying prey items to protective cover before they are consumed will minimize the time spent exposed to predators, whereas feeding at maximum efficiency (staying to eat prey where they are found) will actually increase the time spent exposed to predators. Whether or not there is a conflict between maximizing foraging efficiency and minimizing exposure time, depends upon the travel time to cover relative to the handling time of a prey item; short handling times and/or long travel times are associated with the no-conflict situation, whereas the conflict situation is associated with long handling times and/or short travel times to cover. Free-ranging chickadees foraging at an artificial patch at various distances from cover can distinguish between these two foraging situations. When there is no conflict, they stay and eat at the patch. Their behavior in the conflict situation indicates that they are tradingoff foraging considerations against the risk of predation. When the cost of carrying is low and the benefit gained is high, the chickadees elect to carry items to cover; they tend to stay and eat at the patch when the relative magnitudes of costs and benefits are reversed.     

Food storage in two Late Bronze Age caves of Southern France: palaeoethnobotanical and social implications

This paper presents new archaeobotanical results from two previously studied Late Bronze Age caves situated in Southern France, Balme Gontran and Baume Layrou. At each site a thick black layer, characterised by a very high density of charred seeds, is shown to be composed of the remains of burnt crop stores. In Baume Layrou a small proportion of desiccated plant remains was preserved in addition to the bulk of carbonised material. In Balme Gontran, Triticum spelta and Panicum miliaceum predominated and were independently stored. Lens culinaris, Vicia faba and Setaria italica were secondary species of some importance and could have been stored as well. Storage at Baume Layrou was above all composed of hulled Hordeum vulgare, Triticum spelta and P. miliaceum. Other possibly stored species were Triticum aestivum/durum/turgidum, T. dicoccon and V. faba. It seems that most of the crops were grown in pure stands, with the exception of S. italica and Triticum monococcum which may have been mixed in small proportions with common millet and emmer respectively. Crops were stored in ceramic vessels, probably also in bags and wooden containers like baskets. Millet grains were stored in their husks while glume wheats were dehusked. Dehusking before storage does not seem to have been the common practice at the time. It seems moreover rather unsuitable for grain storage in caves. In Baume Layrou a small proportion of cereal kernels had started to germinate, presumably due to the humidity of the cave. We are making the assumption that the caves were not used for usual long term storage but to store food supplies for a small group of people who intended to live here for a short period, perhaps taking refuge during disturbed times. Crops could have been dehusked to reduce the weight and volume of the load to transport to the caves on steep and difficult paths.