Ian P. F. Owens

Extra pair paternity in birds: a review of interspecific variation and adaptive function.

The application of molecular genetic techniques has revolutionized our view of avian mating systems. Contrary to prior expectations, birds are only very rarely sexually monogamous, with ‘extra‐pair offspring’ found in approximately 90% of species. Even among socially monogamous species, over 11% of offspring are, on average, the result of extra‐pair paternity (EPP). Based on over 150 molecular genetic studies of EPP in birds, we review two topical areas: (i) ecological explanations for interspecific variation in the rate of EPP; and (ii) evidence bearing on the adaptive function of EPP. We highlight the remaining challenges of understanding the relative roles of genes and ecology in determining variation between taxa in the rate of extra paternity, and testing for differences between extra‐pair offspring and those sired within‐pair.

Global hotspots of species richness are not congruent with endemism or threat

Biodiversity hotspots have a prominent role in conservation biology, but it remains controversial to what extent different types of hotspot are congruent. Previous studies were unable to provide a general answer because they used a single biodiversity index, were geographically restricted, compared areas of unequal size or did not quantitatively compare hotspot types. Here we use a new global database on the breeding distribution of all known extant bird species to test for congruence across three types of hotspot. We demonstrate that hotspots of species richness, threat and endemism do not show the same geographical distribution. Only 2.5% of hotspot areas are common to all three aspects of diversity, with over 80% of hotspots being idiosyncratic. More generally, there is a surprisingly low overall congruence of biodiversity indices, with any one index explaining less than 24% of variation in the other indices. These results suggest that, even within a single taxonomic class, different mechanisms are responsible for the origin and maintenance of different aspects of diversity. Consequently, the different types of hotspots also vary greatly in their utility as conservation tools.

Costly sexual signals: are carotenoids rare, risky or required?

Theories of animal signalling emphasize the importance of costliness-to be effective, signals must be dependable; to be dependable, signals must carry costs-and carotenoid-based signals are a favoured example. The traditional view that carotenoids are costly because they are scarce still carries weight. However, biomedical research has led to alternative views on costliness, mainly related to beneficial, but also to detrimental, effects of carotenoids. Recent improvements in our understanding of carotenoids suggest that the relative importance of these mechanisms will soon be determined, leading to a fresh outlook on cost-based signalling.

Global distribution and conservation of rare and threatened vertebrates

Global conservation strategies commonly assume that different taxonomic groups show congruent geographical patterns of diversity, and that the distribution of extinction-prone species in one group can therefore act as a surrogate for vulnerable species in other groups when conservation decisions are being made. The validity of these assumptions remains unclear, however, because previous tests have been limited in both geographical and taxonomic extent. Here we use a database on the global distribution of 19,349 living bird, mammal and amphibian species to show that, although the distribution of overall species richness is very similar among these groups, congruence in the distribution of rare and threatened species is markedly lower. Congruence is especially low among the very rarest species. Cross-taxon congruence is also highly scale dependent, being particularly low at the finer spatial resolutions relevant to real protected areas. ‘Hotspots’ of rarity and threat are therefore largely non-overlapping across groups, as are areas chosen to maximize species complementarity. Overall, our results indicate that ‘silver-bullet’ conservation strategies alone will not deliver efficient conservation solutions. Instead, priority areas for biodiversity conservation must be based on high-resolution data from multiple taxa.

SEXUAL DIMORPHISM IN BIRDS : WHY ARE THERE SO MANY DIFFERENT FORMS OF DIMORPHISM?

Variation in the extent of sexual dimorphism among bird species is traditionally attributed to differences in social mating system. However, there are many different forms of dimorphism among birds, and not all of them show an obvious correlation with social mating system. For example, recent work has shown that many highly polygamous species are, in fact, monomorphic, whereas many putatively monogamous species are dimorphic. In this paper we break up sexual dimorphism into subcomponents and then use comparative analyses to examine the pattern of covariation between these subcomponents and various aspects of sexual, social, and parental behaviour. Our first finding is that size dimorphism and plumage–colour dimorphism do not show the same pattern of covariation. Differences in size dimorphism are associated with variation in social mating system and sex differences in parental care, whereas differences in plumage–colour dimorphism are associated with variation in the frequency of extra–bond paternity. These results suggest that size dimorphism is associated with the sort of intrasexual competition described by traditional classifications of social mating system, whereas plumage–colour dimorphism is associated with cryptic female choice. However, when we break up plumage–colour dimorphism according to whether it is due to melanins, carotenoids or structural colours, we find that each category of plumage–colour dimorphism shows a different pattern of covariation. The correlation between overall plumage–colour dimorphism and the rate of extra–bond paternity is due to structural colours, whereas melanin–based dimorphism is associated with sex differences in parental care. The former result is particularly interesting given that new work suggests structural colours are associated with active sexual displays and the reflection of ultraviolet light.

Variation in extinction risk among birds: chance or evolutionary predisposition?

Collar et al. (1994) estimate that of the 9,672 extant species of bird, 1,111 are threatened by extinction. Here, we test whether these threatened species are simply a random sample of birds, or whether there is something about their biology that predisposes them to extinction. We ask three specific questions. First, is extinction risk randomly distributed among families? Second, which families, if any, contain more, or less, threatened species than would be expected by chance? Third, is variation between taxa in extinction risk associated with variation in either body size or fecundity? Extinction risk is not randomly distributed among families. The families which contain significantly more threatened species than expected are the parrots (Psittacidae), pheasants and allies (Phasianidae), albatrosses and allies (Procellariidae), rails (Rallidae), cranes (Gruidae), cracids (Cracidae), megapodes (Megapodidae) and pigeons (Columbidae). The only family which contains significantly fewer threatened species than expected is the woodpeckers (Picidae). Extinction risk is also not distributed randomly with respect to fecundity or body size. Once phylogeny has been controlled for, increases in extinction risk are independently associated with increases in body size and decreases in fecundity. We suggest that this is because low rates of fecundity, which evolved many tens of millions of years ago, predisposed certain lineages to extinction. Low–fecundity populations take longer to recover if they are reduced to small sizes and are, therefore, more likely to go extinct if an external force causes an increase in the rate of mortality, thereby perturbing the natural balance between fecundity and mortality.

Cooperative breeding in birds: a comparative test of the life history hypothesis

In approximately 3.2% of bird species individuals regularly forgo the opportunity to breed independently and instead breed cooperatively with other conspecifics, either as non–reproductive ‘helpers’ or as co–breeders. The traditional explanation for cooperative breeding is that the opportunities for breeding independently are limited owing to peculiar features of the species breeding ecology. However, it has proved remarkably difficult to find any common ecological correlates of cooperative breeding in birds. This difficulty has led to the ‘life history hypothesis’, which suggests that the common feature of cooperatively breeding birds is their great longevity, rather than any particular feature of their breeding ecology. Here, we use a comparative method to test the life history hypothesis by looking for correlations between life history variation and variation in the frequency of cooperative breeding. First, we find that cooperative breeding in birds is not randomly distributed, but concentrated in certain families, thus supporting the idea that there may be a common basis to cooperative breeding in birds. Second, increases in the level of cooperative breeding are strongly associated with decreases in annual adult mortality and modal clutch size. Third, the proportion of cooperatively breeding species per family is correlated with a low family–typical value of annual mortality, suggesting that low mortality predisposes cooperative breeding rather than vice versa. Finally, the low rate of mortality typically found in cooperatively breeding species is associated with increasing sedentariness, lower latitudes, and decreased environmental fluctuation. We suggest that low annual mortality is the key factor that predisposes avian lineages to cooperative breeding, then ecological changes, such as becoming sedentary, further slow population turnover and reduce opportunities for independent breeding. As the traditional explanation suggests, the breeding habitat of cooperatively breeding species is saturated, but this saturation is not owing to any peculiar feature of the breeding ecology of cooperative breeders. Rather, the saturation arises because the local population turnover in these species is unusually slow, as predicted by the life history hypothesis.

Hormonal basis of sexual dimorphism in birds: implications for new theories of sexual selection

It is widely assumed that the development of male secondary sexual traits in birds and mammals is testosterone-dependent. In birds, however, masculinity has dual origins. Male-type behaviour and morphology, such as spurs and wattles, are usually testosterone-dependent. However, showy male-type plumage is, generally, the neutral state of development. For example, castrating a peacock has no effect on his elaborate plumage whereas ovariectomizing a peahen causes her to develop showy male-type plumage. The surprising relationships between dimorphism and gonadal steroids in birds have important consequences for the current debate concerning the evolution of biological signals and, in particular, the immunocompetence-handicap principle.

Genetic consequences of sequential founder events by an island-colonizing bird

The importance of founder events in promoting evolutionary changes on islands has been a subject of long-running controversy. Resolution of this debate has been hindered by a lack of empirical evidence from naturally founded island populations. Here we undertake a genetic analysis of a series of historically documented, natural colonization events by the silvereye species-complex (Zosterops lateralis), a group used to illustrate the process of island colonization in the original founder effect model. Our results indicate that single founder events do not affect levels of heterozygosity or allelic diversity, nor do they result in immediate genetic differentiation between populations. Instead, four to five successive founder events are required before indices of diversity and divergence approach that seen in evolutionarily old forms. A Bayesian analysis based on computer simulation allows inferences to be made on the number of effective founders and indicates that founder effects are weak because island populations are established from relatively large flocks. Indeed, statistical support for a founder event model was not significantly higher than for a gradual-drift model for all recently colonized islands. Taken together, these results suggest that single colonization events in this species complex are rarely accompanied by severe founder effects, and multiple founder events and/or long-term genetic drift have been of greater consequence for neutral genetic diversity.

Environmental determination of a sexually selected trait

Models of sexual selection usually assume that variation in the expression of sexual ornaments is determined largely by genetic, rather than environmental, factors. However, empirical support for this assumption comes from studies of species with little parental care,, in which the influence of environmental factors may be limited,, and from studies of just two species, with parental care, in both of which heritability estimates vary hugely between years or populations,. In the remaining studies of species with parental care, it is not known whether resemblance in sexual ornamentation between relatives was due to shared genes or shared patterns of care,,. Here we use cross-fostering experiments in house sparrows, Passer domesticus, to examine the relative roles of these effects. We demonstrate that, although sons resemble their fathers with respect to sexual ornamentation, this resemblance is mainly due to post-hatching environmental effects rather than shared genes. We also show that sons hatching early in the year have the largest ornaments. These results support models that emphasize the importance of environmental sources of variation, such as direct paternal effects,,, on the expression of sexual ornaments, and agree with the general observation that sexually selected traits tend to be condition dependent. We urge the incorporation of gene–environment interactions into future models of sexual selection.