Low frequency of extra-pair paternity in two colonies of the socially monogamous short-tailed shearwater Puffinus tenuirostris

The short-tailed shearwater is a colonially nesting, socially monogamous seabird. Little is known about mate fidelity and breeding behaviour in this species because breeding birds are nocturnal on land and spend much of their time within subterranean nesting burrows. Colonial breeding and extended sperm storage create opportunities for extra-pair copulations which may form a significant component of the mating strategy in this species. Multilocus DNA fingerprinting was used to examine the genetic relationship between nestlings and the male and female nest attendants in 83 burrows from two distinct breeding colonies. Genetic analyses identified nine nestlings, approximately equally distributed between the two colonies, that were not related to the attendant pair male in those burrows, implying extra-pair paternity through extra-pair copulations. These results are used retrospectively to discuss the characteristics of extra-pair copulations and extra-pair fertilizations and the implications for estimates of life-time reproductive success in the short-tailed shearwater.     

Resolving genetic relationships with microsatellite markers: a parentage testing system for the swallow Hirundo rustica

Eight polymorphic microsatellite markers from the swallow were isolated and characterized. Extraordinary variability was revealed at the HrU6 locus with 45 different alleles scored among 46 unrelated individuals. The probability that the same genotype combination would occur in two random and unrelated individuals at six selected loci was as low as 1.3 × 10^-8 and the combined exclusion probability was 0.9996. Stable Mendelian inheritance was observed in about 1000 meioses. No significant linkage was revealed and for almost all combinations of marker-pairs, linkage closer than 5 cM could be excluded. At two loci, null (nonamplifying) alleles were encountered. Thirteen (30%) extra-pair offspring were identified in 5 (56%) broods when applying the marker set on a nearly complete swallow colony. We were able to identify a single male from the other families in the colony as the most likely father for nine of the 13 extra-pair offspring.     

Sperm competition in tropical versus temperate zone birds

Sperm competition represents an important component of post-copulatory sexual selection. It has been argued that the level of sperm competition declines in birds towards the equator. However, to date, sperm competition estimates have been available mainly for avian species inhabiting the northern temperate zone. Here we apply a novel approach, using the coefficient of between-male variation (CV_bm) in sperm size as an index for sperm competition risk, in a comparative analysis of 31 Afrotropical and 99 northern temperate zone passerine species. We found no difference in sperm competition risk between the two groups, nor any relationship with migration distance. However, a multivariate model indicated that sperm competition risk was highest in species with a combination of low body mass and few eggs per clutch. The effect of clutch size was most pronounced in tropical species, which indicates that sperm competition risk in tropical and temperate species is differently associated with particular life-history traits. Although tropical species had lower sperm competition risk than temperate zone species for overlapping clutch sizes, the idea of a generally reduced risk of sperm competition in tropical birds was not supported by our analysis.     

The couple that sings together stays together: duetting,aggression and extra-pair paternity in a promiscuous bird species

When individuals mate outside the pair bond, males should employ behaviours such as aggression or vocal displays (e.g. duetting) that help assure paternity of the offspring they care for. We tested whether male paternity was associated with aggression or duetting in the red-backed fairy-wren, a species exhibiting high rates of extra-pair paternity. During simulated territorial intrusions, aggression and duetting were variable among and repeatable within males, suggesting behavioural consistency of individuals. Males with quicker and stronger duet responses were cuckolded less often than males with slower and weaker responses. In contrast, physical aggression was not correlated with male paternity. These results suggest that either acoustic mate guarding or male–female vocal negotiations via duetting lead to increased paternity assurance, whereas physical aggression does not.     

What is behind the variation in mate quality dependent sex ratio adjustment? – A meta‐analysis

Theory predicts that parents adjust the sex ratio of their brood to the sexually selected traits of their mate because the reproductive success of sons may be more dependent on inherited paternal attractiveness than that of daughters. Empirical studies vary in terms of whether they support the theory, and this variation has often been regarded as evidence against sex ratio adjustment or has been ascribed to methodological differences. Applying phylogenetic meta‐analyses, we aimed to find biological explanations for the variation observed in songbirds. In particular, we tested the role extra‐pair paternity, because infidelity occurs in the majority of these species and may reduce the adaptive value of adjusting brood sex ratio to the phenotype of the social mate. However, we found that the variation in effect sizes was unrelated to the proportion of extra‐pair paternity. Thus future studies should consider that mate quality dependent sex ratio adjustment may be driven by direct (material) rather than indirect (genetic) benefits. We also tested if the effect sizes are influenced by whether the focal male trait is indeed under sexual selection as it is assumed by the sex allocation theory. We found that for male traits with proven role in sexual selection, effect sizes significantly differed from the null expectation of random production of sons and daughters. For male traits with only presumed sexual role in sexual selection, the deviation from the null expectation was less convincing, and the effect sizes were significantly smaller. This result indicates that studies that neglect the assumptions of the hypotheses concerned, may lead to the underestimation of the mean effect size and, eventually, false conclusions.     

Molecular insight into patterns of colony composition and paternity in the common mole-rat Cryptomys hottentotus hottentotus

We report the discovery of intraspecific variation in both colony composition and patterns of paternity in two populations of the social common mole-rat Cryptomys hottentotus hottentotus. These two populations represent the mesic and arid habitat extremes of the species' broad ecological range in South Africa. Until recently colonies of the common mole-rat were thought to consist of familial groups whereby all colony members were the offspring of a monogamous reproductive pair. The remaining colony members were thought to forego reproduction until both social and ecological conditions favoured dispersal and opportunities for independent outbreeding. Results from genetic assignment tests using microsatellite markers indicate that while colony composition is dominated by familial groups, colonies within both populations included both adult and subadult foreign conspecifics. Analysis of parentage reveals that the social organization of C. h. hottentotus is not that of strict monogamy; paternity of offspring was not assigned consistently to the largest, most dominant male within the colony. Moreover, a number of significantly smaller males were found to sire offspring, suggesting a sneak-mating strategy by subordinate within-colony males. Extra-colony extra-pair paternity (ECP) was also found to characterize C. h. hottentotus colonies, occurring with similar frequencies in both habitats. Both dominant established breeding males and subordinate males were identified as siring young in nonsource colonies. Furthermore, established breeding males were found to sire extra-colony young in the same season as siring young within their source colonies. We discuss the significance of these results within the context of the divergent ecological regimes characterizing the two sites and observe that our results revisit the accuracy of using behavioural and morphological characters, which have structured the basis of our understanding of the behavioural ecology of this species, as indicators of breeding status in mark-recapture studies.     

Extra-pair paternity in waved albatrosses

We estimated the rate of extra‐pair fertilizations (EPFs) in waved albatrosses (Phoebastria irrorata) on Isla Española, Galápagos, Ecuador, using multilocus minisatellite DNA fingerprinting. Waved albatrosses are socially monogamous, long‐lived seabirds whose main population is on Española. Aggressive extra‐pair copulation (EPC) attempts have been observed in the breeding colony during the days preceding egg‐laying. Our genetic analyses of 16 families (single chicks and their attending parents) revealed evidence of EPFs in four families. In all cases males were the excluded parent. These data suggest that waved albatrosses have an unusually high rate of EPF relative to taxa with similar life histories. Future behavioural observations will determine the extent to which forced vs. unforced EPCs contribute to this high EPF rate.     

Female sexual receptivity after partial copulations in the wolf spider (Schizocosa malitiosa)

Mated females of the wolf spider Schizocosa malitiosa are frequently refractory to further copulations. Copulation consists of two successive behavioural patterns: pattern I (PI) and pattern II (PII). During PI males make multiple consecutive insertions with each palp, and in PII males alternate the use of palps after each insertion until dismounting. As both patterns are inseminatory, another function – such as generating female reluctance – is suggested for this complex behaviour. Here we test experimentally whether female spiders mated only with PI, or only with PII, are reluctant to re-mate. Each copulating male was interrupted immediately after the end of PI (phase A), and the same male was instantly exposed to a second virgin female. After re-mounting, the male initially performed a brief recapitulation of PI followed by PII (both together considered as phase B). For testing female sexual reluctance, females were exposed to a second male 3 days after the partial mating, but whenever the mounting occurred it was interrupted. Females were raised and progeny was counted. Phase-A-mated females were more reluctant to re-mating than phase-B-mated females, but their respective progeny was similar. We suggest that female reluctance is caused by sperm-associated substances transferred during PI.     

Comparison of the mating systems and breeding behavior of a resident and a migratory tropical flycatcher

ABSTRACT.   Little is known about the genetic mating systems of tropical passerines and how they vary among species. We studied the Lesser Elaenia ( Elaenia chiriquensis ) and the Yellow-bellied Elaenia ( E. flavogaster ) near Gamboa, Panama. These species breed in the same habitat, but Lesser Elaenias are intratropical migrants with seasonal territories and Yellow-bellied Elaenias are permanent residents that remain paired and defend territories throughout the year. Lesser Elaenias exhibited greater breeding synchrony (15–18 %) than Yellow-bellied Elaenias (9–10%). For Lesser Elaenias, 10 of 15 (67%) nests contained extra-pair young and 14 of 38 (37%) young resulted from extra-pair fertilizations (EPFs). In contrast, only one extra-pair nestling (4%, N = 24 nestlings) was found in 13 Yellow-bellied Elaenia nests. Neither species exhibited strong mate guarding. The higher rate of EPFs in Lesser Elaenias is consistent with the hypothesis that year-round territorial tropical passerines with low breeding synchrony have little or no extra-pair behavior compared with species that breed seasonally. Although the low singing rates of Lesser Elaenias (7 songs/h) suggest that this not an important cue for female extra-pair mate choice, the role of conspicuous male dawn song remains to be investigated. Further studies of tropical passerines are needed to help disentangle the effects of synchrony, density, and other ecological and behavioral factors that have influenced the evolution of extra-pair mating systems in passerines.     

How do misassigned paternities affect the estimation of heritability in the wild?

Studies of birds have recently played an important role in the increasing success of quantitative genetics applied to natural populations. However, these studies mostly base their pedigree relationships on social information, despite the known widespread genetic polygamy in avian species. Here, we study the influence of misassigned paternities, combined with the effect of pedigree size and depth, on the estimation of heritability. First, we compute simulations of a polygenic trait for two levels of heritability (0.1 and 0.4), several extra-pair paternity rates (ranging from 5% to 40%), and varying sample sizes (20, 50 and 100 broods) or pedigree depth (2 or 4 generations). We compare heritability estimates from the social and the genetic pedigree, running a restricted maximum-likelihood 'animal model'. Social pedigree underestimates heritability by an average of 0-17% for 5-20% extra-pair paternities and by up to 18% for 40% extra-pair paternities and a heritability of 0.4. Second, we identifyied extra-pair offspring using microsatellite loci in two populations of blue tits (Parus caeruleus) showing high levels of extra-pair paternities (15% and 25% of extra-pair offspring). We compare heritabilities of tarsus length and body mass estimated with pedigrees of increasing accuracy. These analyses suggest that the bias induced by misassigned paternities on heritability estimation depends on the level of heritability and the rate of paternity error. Typical rates of extra-pair paternities in birds (around 20% of offspring) should result in an underestimation of heritability of less than 15% when estimated over a minimum of 100 broods.