Laurene M. Ratcliffe

Tropical Winter Habitat Limits Reproductive Success on the Temperate Breeding Grounds in a Migratory Bird

Identifying the factors that control population dynamics in migratory animals has been constrained by our inability to track individuals throughout the annual cycle. Using stable carbon isotopes, we show that the reproductive success of a long‐distance migratory bird is influenced by the quality of habitat located thousands of kilometres away on tropical wintering grounds. For male American redstarts (Setophaga ruticilla), winter habitat quality influenced arrival date on the breeding grounds, which in turn affected key variables associated with reproduction, including the number of young fledged. Based on a winter‐habitat model, females occupying high‐quality winter habitat were predicted to produce more than two additional young and to fledge offspring up to a month earlier compared with females wintering in poor‐quality habitat. Differences of this magnitude are highly important considering redstarts are single brooded, lay clutches of only three to five eggs and spend only two‐and‐a‐half months on the breeding grounds. Results from this study indicate the importance of understanding how periods of the annual cycle interact for migratory animals. Continued loss of tropical wintering habitat could have negative effects on migratory populations in the following breeding season, minimizing density‐dependent effects on the breeding grounds and leading to further population declines. If conservation efforts are to be successful, strategies must incorporate measures to protect all the habitats used during the entire annual cycle of migratory animals.

Do female black-capped chickadees prefer high-ranking males as extra-pair partners ?

Abstract Previous studies have shown that some female black-capped chickadees (Poecile atricapillus) solicit copulations from males that rank higher in winter flocks than their social mates, and extra-pair paternity in nests occurs commonly enough to be considered a potential female mating tactic. This study uses blood samples collected in 1992–1995 from 58 families of black-capped chickadees to test whether females with extra-pair offspring have chosen extra-pair sires higher in social rank than their mates. Paternity was assessed with multilocus DNA fingerprinting in 1992–1994 nests and with microsatellite and single-locus minisatellite DNA typing in 1995 nests. Seventeen of 58 nests (29.3%) contained young genetically mismatched with their social father. In 11 of 15 cases where the identity of the extra-pair male was known, the extra-pair male was dominant to the social father. Using data from 29 nests located in 1994 and 1995 for which we had the most data on relative ranks of males, high-ranking males had greater realized reproductive success than low-ranking males as a result of extra-pair fertilizations. There was no significant difference between the number of nests containing extra-pair young of females mated to low-ranked versus high-ranked males. Two nests in 1995 contained young either genetically mismatched with both social parents (intraspecific brood parasitism) or, in one nest, genetically mismatched with the social mother but not the social father (quasi-parasitism). The implications of female strategies acquiring genetic benefits through extra-pair copulations are discussed.

Design of Playback Experiments: The Thornbridge Hall NATO ARW Consensus

Playback is an experimental technique commonly used to investigate the significance of signals in animal communication systems. It involves replaying recordings of naturally occurring or synthesised signals to animals and noting their response. Playback has made a major contribution to our understanding of animal communication, but like any other technique, it has its limitations and constraints.

Pitch shifts and song structure indicate male quality in the dawn chorus of black-capped chickadees

The fee-bee song of male black-capped chickadees (Poecile atricapillus) is considered a single-type song that singers transpose up and down a continuous frequency range. While the ability to shift song pitch in this species provides a mechanism for song matching as an aversive signal in male-male territorial song contests, the functional significance of this behaviour during the “solo” performances of males during the dawn chorus is unclear. We analysed the dawn chorus songs and singing behaviour of males whose winter-flock dominance status we determined. We used correlation analysis to show that pitch shifts were accompanied by changes to other fine structural characteristics in song, including temporal and relative amplitude parameters. We also found that songs of socially dominant males and songs of their most subordinate flockmates could be distinguished using these methods by the way they performed a between-note frequency measure accompanying pitch shifts. That is, a ratio measure of the internote frequency interval remained constant for songs of high-ranking birds despite changes in absolute pitch, while low-ranking males sang a smaller ratio as they shifted to higher absolute pitches. These findings identify previously unrecognised variation in the songs of black-capped chickadees. More importantly, they indicate a mechanism by which pitch shifting during the dawn chorus of black-capped chickadees could provide a reliable indicator of relative male quality.

Non-breeding season events influence sexual selection in a long-distance migratory bird

The study of sexual selection has traditionally focused on events and behaviours immediately surrounding copulation. In this study, we examine whether carry-over effects from the non-breeding season can influence the process of sexual selection in a long-distance migratory bird, the American redstart (Setophaga ruticilla). Previous work on American redstarts demonstrated that overwintering in a high-quality habitat influences spring departure dates from the wintering grounds, advances arrival dates on the breeding grounds and increases apparent reproductive success. We show that the mixed-mating strategy of American redstarts compounds the benefits of overwintering in high-quality winter habitats. Males arriving to breed in Canada from high-quality winter habitats arrive earlier than males from poor-quality habitats, resulting in a lower probability of paternity loss, a higher probability of achieving polygyny and ultimately higher realized reproductive success. Such results suggest that the process of sexual selection may be influenced by events interacting throughout the annual cycle.

Achromatic color variation in black-capped chickadees, Poecile atricapilla : black and white signals of sex and rank

Sexual dichromatism and phenotypic variation in elaborate male traits are common products of sexual selection. The spectral properties of carotenoid and structurally-based plumage colors and the patch sizes of melanin-based plumage colors have received considerable attention as sexual signals in birds. However, the importance of variation in achromatic plumage colors (white, gray and black) remains virtually unexplored, despite their widespread occurrence. We investigated a potential signal function of the achromatic black and white plumage of black-capped chickadees (Poecile atricapilla). We captured and color-banded 178 free-living chickadees and assessed winter flock dominance hierarchies by tabulating pairwise interactions at feeders. We recaptured 73 of these birds and measured plumage coloration for six body regions using a reflectance spectrometer and the area of melanin-based plumage patches from standardized photographs. We found extensive individual variation in chickadee plumage traits and considerable sexual dichromatism. Male black-capped chickadees have significantly brighter white plumage than females, larger black patches, and greater plumage contrast between adjacent white and black plumage regions. We also found rank differences in the plumage reflectance of males; high-ranking males, who are preferred by females as both social and extra-pair partners, exhibit significantly darker black plumage and grow their feathers more rapidly than low-ranking males. This variation among individuals reveals a potential signal function for achromatic plumage coloration in birds.

Disruptive sexual selection for plumage coloration in a passerine bird

The theory of sexual selection was developed to explain the evolution of highly exaggerated sexual ornaments. Now supported by vast empirical evidence, sexual selection is generally considered to favour individuals with the most extreme trait expression. Here we describe disruptive selection on a sexual ornament, plumage coloration, in yearling male lazuli buntings (Passerina amoena). In habitats with limited good-quality nesting cover, the dullest and the brightest yearlings were more successful in obtaining high-quality territories, pairing with females and siring offspring, than yearlings with intermediate plumage. This pattern reflects the way that territorial adult males vary levels of aggression to influence the structure of their social neighbourhood. Adult males showed less aggression towards dull yearlings than intermediate and bright ones, permitting the dull yearlings to settle on good territories nearby. Fitness comparisons based on paternity analyses showed that both the adults and dull yearlings benefited genetically from this arrangement, revealing a rare example of sexually selected male–male cooperation.

Absolute and Relative Pitch Production in the Song of the Black-Capped Chickadee

Black-capped Chickadee (Parus atricapillus) song consists of two notes, termed fee and bee. Frequency measures at three key points (at the start and end of fee, and at the start of bee) were obtained from the songs of a large sample of chickadees (n = 151) in the wild. In this sample, 19 birds produced songs shifted downward in frequency as well as their normal songs. Analysis of normal song revealed that fee declines in frequency in a glissando of nearly pure tone, then continues at greatly reduced amplitude at the start of bee; whereas bee, also a nearly pure tone, is always lower in frequency than either the start or end offee. The absolute pitches (frequencies) of these measures vary substantially among birds, but much less within individuals. In contrast, pitch intervals (ratios of higher to lower frequencies) for frequency changes among the three measures are highly invariant among birds. Moreover, chickadees with normal and frequency-shifted songs maintain virtually the same pitch intervals in both. This analysis suggests that the absolute and relative pitch constancies in chickadee song production may provide information for individual and species recognition, respectively.

Female initiated divorce in a monogamous songbird: abandoning mates for males of higher quality

Divorcing a current partner to re-pair with a mate of higher quality may be a strategy to increase reproductive success used by socially monogamous birds. By increasing the availability of males through selective mate removal during the nest building period, we found that female black-capped chickadees, Parus atricapillus, will desert their mates to pair with males of higher social rank, a trait closely associated with resource holding potential in this species. Females from neighbouring territories were more likely to desert their mates for high ranked rather than the low ranked widower males; six of seven high ranking and one of six low ranking males were chosen by the divorcing females. Six of the seven widower males chosen by divorcing females were dominant to the males that the females deserted. Once released, the originally removed females were able to re-establish pairbonds with their mates. Higher social ranks of removed females may have enabled them to exclude lower ranked replacement females, and may reflect a natural constraint on female choice.

MITOCHONDRIAL DNA HOMOGENEITY IN THE PHENOTYPICALLY DIVERSE REDPOLL FINCH COMPLEX (AVES: CARDUELINAE: CARDUELIS FLAMMEA-HORNEMANNI)

Breeding redpoll finches (Aves: Carduelinae) show extensive plumage and size variability and, in many cases, a plumage polymorphism that is not related to age or sex. This has been ascribed to extreme phenotypic variation within a single taxon or to moderate variability within distinct taxa coupled with hybridization. The predominant view favors the recognition of two largely sympatric species: Carduelis flammea, comprised of four well‐marked subspecies—flammea, cabaret, islandica, and rostrata; and C. hornemanni, comprised of two subspecies—hornemanni and exilipes. We studied representative samples of these putative subspecies (except islandica) for variation in mitochondrial DNA (mtDNA). Using 20 informative restriction enzymes that recognized 124 sites (642 base pairs [bp] of sequence or ≈ 3.7% of the molecule), we identified 17 RFLP haplotypes in the 31 individuals surveyed. The haplotypes formed a simple phylogenetic network with most clones diverging by a single site difference from a common haplotype found in almost half of the individuals. Within populations and taxa, levels of mtDNA diversity were similar to those observed in other avian species. The pattern of mtDNA divergence among populations was statistically unrelated to their geographic or traditional taxonomic relationships, and the estimated distance between the two traditionally recognized species was very small relative to those typically observed among avian sister species.