Raleigh J. Robertson

The relationship between fitness and baseline glucocorticoids in a passerine bird

Glucocorticoid (cort) hormones are increasingly applied in studies of free-ranging animals, with elevated baseline cort levels generally assumed to indicate individuals or populations in worse condition and with lower fitness (the Cort-Fitness Hypothesis). The relationship between cort and fitness is rarely validated and studies investigating the cort-fitness relationship often find results inconsistent with the Cort-Fitness Hypothesis. The inconsistency of these studies may result in part from variation in the cort-fitness relationship across life history stages. Here we address the following questions in a two-year study in free-ranging tree swallows (Tachycineta bicolor): (1) Do baseline cort levels correlate with fitness within a life history stage? (2) Does the cort-fitness relationship vary across different life history stages? (3) Does the cort-fitness relationship vary across life history stages within an individual? (4) Does reproductive effort influence cort levels, and do cort levels influence reproductive effort? We measured baseline cort and fitness components in female birds of known breeding stages. We find correlations between baseline cort levels and fitness within some life history stages, but the relationship shifts from negative during early breeding to positive during late breeding, even within the same individuals. A positive relationship between baseline cort and fitness components during the nestling period suggests that reproductive investment may elicit higher cort levels that feedback to reallocate more effort to reproduction during critical periods of nestling provisioning. Our findings provide reason to question the Cort-Fitness Hypothesis, and have implications for the application of cort measures in monitoring the condition of populations of conservation concern.

Confidence of Paternity and Male Parental Care

We present a theoretical framework in order to understand how the relationship between male parental care and paternity is dependent on the relationship between male parental care and offspring recruitment. When there is an S-shaped relationship between offspring recruitment and the parental care of a single male, we predict a threshold relationship between male parental care and paternity. Tree swallows, monogamous dunnocks, and red-winged black-birds may follow this pattern. Alternatively, when there is a concave-down relationship between offspring recruitment and male parental care, we predict a continuous (gradual) decline in male parental care in response to decreasing paternity. Studies of noisy miners and polyandrous dunnocks suggest this pattern. Our model illustrates that the relationship between male parental care and paternity is more complex than assumed previously. A predicted adjustment of male parental care in response to paternity must be examined in relation to the effect of male parental care on offspring recruitment and the effect of alternative activities on male fitness

Nest-site characteristics, reproductive success and cavity availability for tree swallows breeding in natural cavities

Etude de 10 caracteristiques des cavites naturelles et de leur influence sur le succes reproducteur de Tachycineta bicolor.Comparaison des sites de nidification de T.b. avec ceux des autres especes et avec des cavites inoccupees dans des habitats similaires et discussion des facteurs influencant la disponibilite en cavites convenables

Female control of extra-pair fertilization in tree swallows

SummaryIn a Canadian population of tree swallows, Tachycineta bicolor, DNA fingerprinting has previously shown that half of all broods contain many offspring resulting from extra-pair copulations (EPCs), whereas the other half contain only legitimate offspring. This bimodal pattern of extra-pair paternity might be due to variation in the effectiveness of male paternity guards, variation in female ability to resist EPCs, and/or variation in female pursuit of EPCs. Here we report experimental evidence for female control of copulations and fertilizations and the occurrence of two alternative copulation strategies among females in this population. Ten paired male tree swallows were removed on the day their mates laid the first egg. Replacement males took over the nestbox within 0.5–23 h and attempted to copulate with the widowed female. Assuming that eggs were fertilized approximately 24 h prior to laying, the first two eggs were fertilized before the male was removed, while the third and subsequent eggs could potentially be fertilized by the replacement male. Fingerprinting revealed that the first two eggs were sired by the resident males in five nests and by extra-pair males in the remaining five nests. The widows that had been faithful to their initially chosen mate rejected copulation attempts by the replacement male until most of the eggs had been laid. Consequently, nearly all eggs laid by these females were sired by the original male. The widows that had been unfaithful prior to male removal copulated sooner with the replacement male than females that were faithful to their mate. However, these replacement males also had a very low fertilization success; most eggs were sired by males that were not associated with the nest. This is consistent with the situation in non-experimental nests where unfaithful females copulate with their mate at the same rate as faithful females, yet unfaithful females have a majority of offspring sired by extra-pair males. We conclude that fertilization patterns to a large extent are determined by the female through active selection and rejection of copulation partners, though our results also allow some speculation that females have control over sperm competition. Female copulation tactics are probably determined some currently unknown fitness benefits of having the offspring sired by particular males.

Extra-pair paternity in tree swallows: why do females mate with more than one male?

Recent studies of monogamous tree swallows (Tachycineta bicolor) suggest that females may receive some type of genetic benefit from extra-pair fertilizations. In this study we attempted to determine what type of genetic benefits might be gained by females. We compared numerous morphological and behavioral traits (Table 1) of every male nesting on one grid of nest-boxes (n = 23) to determine what male traits were correlated with male success at gaining extra-pair fertilizations. DNA fingerprinting revealed an increase in the level of extra-pair paternity from the previous year (50% of broods contained extra-pair young in 1990 vs. 87% of broods in 1991), but no significant correlates of paternity. We found six extra-pair fathers at seven nests (20 nests had extra-pair young). The traits of these extra-pair males did not differ from those of the males they cuckolded. We discuss several reasons for this lack of difference, but argue that our results are not inconsistent with females choosing extra-pair males to enhance the genotypic quality of their offspring. Despite a complete search of the nest-box grid for extra-pair fathers, we were able to explain the paternity of just 21% (13/63) of all extra-pair young. This suggests that extra-pair fathers were either residents off our study grid or non-territorial floaters. Tree swallows are quite mobile and spend only part of the day at their nest prior to laying. In addition, we rarely see swallows visiting other grids of nest-boxes. Therefore, we suggest that most extra-pair copulations occur at some unknown location, possibly at a feeding or roosting area where females may be able to choose from many more potential extra-pair fathers than at their nest-site.

Life-history variation predicts the effects of demographic stochasticity on avian population dynamics

Comparative analyses of avian population fluctuations have shown large interspecific differences in population variability that have been difficult to relate to variation in general ecological characteristics. Here we show that interspecific variation in demographic stochasticity, caused by random variation among individuals in their fitness contributions, can be predicted from a knowledge of the species’ position along a “slow‐fast” gradient of life‐history variation, ranging from high reproductive species with short life expectancy at one end to species that often produce a single offspring but survive well at the other end of the continuum. The demographic stochasticity decreased with adult survival rate, age at maturity, and generation time or the position of the species toward the slow end of the slow‐fast life‐history gradient. This relationship between life‐history characteristics and demographic stochasticity was related to interspecific differences in the variation among females in recruitment as well as to differences in the individual variation in survival. Because reproductive decisions in birds are often subject to strong natural selection, our results provide strong evidence for adaptive modifications of reproductive investment through life‐history evolution of the influence of stochastic variation on avian population dynamics.

Equality of feeding roles and the maintenance of monogamy in tree swallows

SummaryWe investigated the division of labor in male and female tree swallows by measuring the rate of food delivery to nestlings at 36 nests. By observing natural nests and performing brood manipulations we found that males and females divided the feeding duties about equally and responded similarly to changes in brood size and age. Feeding rate was most highly correlated with brood mass. Manipulation and removal experiments showed that increased feeding rates could be elicited, but only for limited periods of time. Male and female tree swallows could only partially compensate in feeding nestlings when mates were removed. This, along with the higher mortality in enlarged broods and in those raised by single parents, indicates that both male and female are required to raise an entire brood to fledging. We argue that this requirement contributes to the absence of mate guarding and the maintenance of monogamy in the tree swallow.

Spatial and temporal multi-species nesting aggregations in birds as anti-parasite and anti-predator defenses

Summary1.The purpose of this study was to determine whether the yellow warbler (Dendroica petechia) derives any benefits from nesting in close proximity to its own and other species in reducing predator pressure or the incidence of brood parasitism by the brown-headed cowbird (Molothrus ater).2.Yellow warblers nesting synchronously with their own and with neighbouring species resulted in a proportionate reduction in the number of nests preyed upon, suggesting a ‘swamping effect’ of the local predators. A significantly lower incidence of predation occurred at yellow warbler nests which were inside a gray catbird (Dumetella carolinensis) response range. This was thought to be the result of yellow warblers taking advantage of the catbirds nest-guarding behaviour and aggressive responses to predators.3.Yellow warblers nesting synchronously with other yellow warblers were subject to a proportionately lower incidence of brood parasitism than asynchronous nests, suggesting a ‘swamping effect’ on the cowbird. A significantly lower incidence of brood parasitism occurred at yellow warbler nests which were in a red-winged blackbird (Agelaius phoeniceus) response range. The yellow warbler was thought to be taking advantage of the aggressive response of red-winged blackbirds to cowbirds.4.Overall, yellow warblers which nested within the response range of red-winged blackbirds or gray catbirds had significantly higher nest success than other yellow warblers. This suggests that selection for defence against predators and cowbirds may lead to multi-species aggregations.

Intraspecific Variation in Sperm Length is Negatively Related to Sperm Competition in Passerine Birds

Abstract Spermatozoa are among the most diversified cells in the animal kingdom, but the underlying evolutionary forces affecting intraspecific variation in sperm morphology are poorly understood. It has been hypothesized that sperm competition is a potent selection pressure on sperm variation within species. Here, we examine intraspecific variation in total sperm length of 22 wild passerine bird species (21 genera, 11 families) in relation to the risk of sperm competition, as expressed by the frequency of extrapair paternity and relative testis size. We demonstrate, by using phylogenetic comparative methods, that between-male variation in sperm length within species is closely and negatively linked to the risk of sperm competition. This relationship was even stronger when only considering species in which data on sperm length and extrapair paternity originated from the same populations. Intramale variation in sperm length within species was also negatively, although nonsignificantly, related to sperm competition risk. Our findings suggest that postcopulatory sexual selection is a powerful evolutionary force reducing the intraspecific phenotypic variation in sperm-size traits, potentially driving the diversification of sperm morphology across populations and species.

COMPARATIVE EVIDENCE FOR THE EVOLUTION OF SPERM SWIMMING SPEED BY SPERM COMPETITION AND FEMALE SPERM STORAGE DURATION IN PASSERINE BIRDS

Sperm swimming speed is an important determinant of male fertility and sperm competitiveness. Despite its fundamental biological importance, the underlying evolutionary processes affecting this male reproductive trait are poorly understood. Using a comparative approach in a phylogenetic framework, we tested the predictions that sperm swim faster with (1) increased risk of sperm competition, (2) shorter duration of female sperm storage, and (3) increased sperm length. We recorded sperm swimming speed in 42 North American and European free-living passerine bird species, representing 35 genera and 16 families. We found that sperm swimming speed was positively related to the frequency of extrapair paternity (a proxy for the risk of sperm competition) and negatively associated with clutch size (a proxy for the duration of female sperm storage). Sperm swimming speed was unrelated to sperm length, although sperm length also increased with the frequency of extrapair paternity. These results suggest that sperm swimming speed and sperm length are not closely associated traits and evolve independently in response to sperm competition in passerine birds. Our findings emphasize the significance of both sperm competition and female sperm storage duration as evolutionary forces driving sperm swimming speed.