The effects of anthropogenic alteration of nesting habitat on rates of extra‐pair fertilization and intraspecific brood parasitism in Canada Geese Branta canadensis

Parentage studies have shown that alternative reproductive strategies are widespread in many avian taxa that were once thought to be monogamous. Recent anthropogenically mediated habitat change may have disrupted ecological factors, such as breeding density, which have given rise to inter‐ and intraspecific variation in the frequency of extra‐pair fertilization (EPF) and intraspecific brood parasitism (IBP). We used genetic analyses to quantify the incidence of alternative reproductive strategies exhibited within clutches of Canada Geese Branta canadensis maxima nesting in high‐ and low‐density situations in and around urban areas in southern Michigan, USA. We tested the hypothesis that high nesting density would increase the frequency of EPF and IBP. There were no significant differences in rates of EPF and IBP clutches (14 and 26% of clutches, respectively) from nests in high‐density (21.7% EPF, 21.7% IBP) vs. low‐density (5.3% EPF, 31.6% IBP) areas, although high‐density sites had a fourfold higher rate of EPF. Rates of EPF and IBP in high‐density urban areas in Michigan were comparable to rates observed in other species nesting under different ecological conditions. Levels of relatedness between host and parasitic females were higher than expected by chance, suggesting that related females are more tolerant of one another and that host females could gain inclusive fitness benefits from rearing parasitic offspring. Our study highlights the importance of understanding the different costs and benefits associated with alternative behavioural repertoires that may vary as habitats and associated selection pressures are increasingly modified by human activities.     

Phylogenetic effects on functional traits and life history strategies of Australian freshwater fish

Understanding the biogeographic and phylogenetic basis to interspecific differences in species’ functional traits is a central goal of evolutionary biology and community ecology. We quantify the extent of phylogenetic influence on functional traits and life‐history strategies of Australian freshwater fish to highlight intercontinental differences as a result of Australia's unique biogeographic and evolutionary history. We assembled data on life history, morphological and ecological traits from published sources for 194 Australian freshwater species. Interspecific variation among species could be described by a specialist–generalist gradient of variation in life‐history strategies associated with spawning frequency, fecundity and spawning migration. In general, Australian fish showed an affinity for life‐history strategies that maximise fitness in hydrologically unpredictable environments. We also observed differences in trait lability between and within life history, morphological and ecological traits where in general morphological and ecological traits were more labile. Our results showed that life‐history strategies are relatively evolutionarily labile and species have potentially evolved or colonised in freshwaters frequently and independently allowing them to maximise population performance in a range of environments. In addition, reproductive guild membership showed strong phylogenetic constraint indicating that evolutionary history is an important component influencing the range and distribution of reproductive strategies in extant species assemblages. For Australian freshwater fish, biogeographic and phylogenetic history contribute to broad taxonomic differences in species functional traits, while finer scale ecological processes contribute to interspecific differences in smaller taxonomic units. These results suggest that the lability or phylogenetic relatedness of different functional traits affects their suitability for testing hypothesis surrounding community level responses to environmental change.     

Extra‐pair paternity in birds

Since the first molecular study providing evidence for mating outside the pair bond in birds over 30 years ago, >500 studies have reported rates of extra‐pair paternity (EPP) in >300 bird species. Here, we give a detailed overview of the current literature reporting EPP in birds and highlight the sampling biases and patterns in the data set with respect to taxonomy, avian phylogeny and global regions, knowledge of which will be crucial for correct interpretation of results in future comparative studies. Subsequently, we use this comprehensive dataset to simultaneously test the role of several ecological and life history variables. We do not find clear evidence that variation in EPP across socially monogamous species can be explained by latitude, density (coloniality), migration, generation length, genetic structuring (dispersal distance), or climatic variability, after accounting for phylogeny. These results contrast previous studies, most likely due to the large heterogeneity within species in both EPP and the predictor of interest, indicating that using species averages might be unreliable. Despite the absence of broadscale ecological drivers in explaining interspecific variation in EPP, we suggest that certain behaviours and ecological variables might facilitate or constrain EPP, as indicated by our finding that EPP was negatively associated with latitude within noncolonial species, suggesting a role of breeding synchrony. Thus, rather than focussing on general explanations for variation in EPP across all species, a future focus should be on how various aspects of ecology or life history might have driven variation in EPP among groups of species or populations of the same species. Hence, we argue that variation in EPP can be partly explained when taking the right perspective. This comprehensive overview, and particularly the dataset provided herein will create a foundation for further studies.     

High fidelity: extra‐pair fertilisations in eight Charadrius plover species are not associated with parental relatedness or social mating system

Extra‐pair paternity is a common reproductive strategy in many bird species. However, it remains unclear why extra‐pair paternity occurs and why it varies among species and populations. Plovers (Charadrius spp.) exhibit considerable variation in reproductive behaviour and ecology, making them excellent models to investigate the evolution of social and genetic mating systems. We investigated inter‐ and intra‐specific patterns of extra‐pair parentage and evaluated three major hypotheses explaining extra‐pair paternity using a comparative approach based on the microsatellite genotypes of 2049 individuals from 510 plover families sampled from twelve populations that constituted eight species. Extra‐pair paternity rates were very low (0 to 4.1% of chicks per population). No evidence was found in support of the sexual conflict or genetic compatibility hypotheses, and there was no seasonal pattern of extra‐pair paternity (EPP). The low prevalence of EPP is consistent with a number of alternative hypotheses, including the parental investment hypothesis, which suggests that high contribution to care by males restricts female plovers from engaging in extra‐pair copulations. Further studies are needed to critically test the importance of this hypothesis for mate choice in plovers.     

Genetic monogamy in two long‐lived New Zealand passerines

High rates of extra‐pair paternity (EPP) can be relatively common in passerines whereas low rates or absence of EPP are often associated with taxa that are long‐lived and exhibit obligatory paternal care. We examined EPP in an under‐represented category: passerine species with relatively long life spans (or low annual mortality rates). Specifically, we studied EPP in New Zealand saddlebacks Philesturnus carunculatus and robins Petroica australis, two species with unusually low annual mortality rates (6.5–11% and 10–20% respectively). No EPP (0%) was detected in saddlebacks (39 pairs, 202 offspring) and only one case of EPP (1.9%) was detected in robins (54 pairs, 198 offspring). Genetic monogamy in these passerine species supports the hypothesis that low annual mortality rates play an important role in explaining variation in rates of EPP across species.     

Multiple hypotheses explain variation in extra‐pair paternity at different levels in a single bird family

Extra‐pair paternity (EPP), where offspring are sired by a male other than the social male, varies enormously both within and among species. Trying to explain this variation has proved difficult because the majority of the interspecific variation is phylogenetically based. Ideally, variation in EPP should be investigated in closely related species, but clades with sufficient variation are rare. We present a comprehensive multifactorial test to explain variation in EPP among individuals in 20 populations of nine species over 89 years from a single bird family (Maluridae). Females had higher EPP in the presence of more helpers, more neighbours or if paired incestuously. Furthermore, higher EPP occurred in years with many incestuous pairs, populations with many helpers and species with high male density or in which males provide less care. Altogether, these variables accounted for 48% of the total and 89% of the interspecific and interpopulation variation in EPP. These findings indicate why consistent patterns in EPP have been so challenging to detect and suggest that a single predictor is unlikely to account for the enormous variation in EPP across levels of analysis. Nevertheless, it also shows that existing hypotheses can explain the variation in EPP well and that the density of males in particular is a good predictor to explain variation in EPP among species when a large part of the confounding effect of phylogeny is excluded.     

Breeding synchrony and extrapair paternity in a species with alternative reproductive strategies

Breeding synchrony may affect the tradeoff between pursuing multiple mates and avoiding paternity loss, translating into differences in the rate of extrapair paternity (EPP). However, diverse empirical relationships between breeding synchrony and EPP remain challenging to explain. We examined whether the relationship between breeding synchrony and EPP varied with male morph, age, body size, or breeding density in the white‐throated sparrow Zonotrichia albicollis. In this species, males of two genetically determined morphs pursue alternative mating strategies. Breeding synchrony positively correlated with EPP within polygamous white morph males, which have high rates of EPP and cuckoldry, but was unrelated to EPP within tan morph males, which prioritize mate guarding and paternal care. As previously reported, males that gained EPP were primarily white males. Males gained EPP more often than expected by chance during their mate's fertile period and on neighboring territories. Since extrapair copulation appears primarily male‐driven in this species, these results indicate that white males focus extra‐pair mating effort during periods of high synchrony and during their mates’ fertile periods, even at the expense of paternity loss within their own nests. Breeding density, male age, and male size did not modify the relationship between breeding synchrony and EPP. However, older white males had higher cuckoldry rates, perhaps reflecting declines in performance associated with senescence. Results suggest that, even within species, mating strategy may modify how breeding synchrony affects rates of EPP, with positive relationships manifest only within subsets of individuals that pursue a strategy of polygyny at the expense of paternity loss.     

Extra-pair paternity does not result in differential sexual selection in the mutually ornamented black swan (Cygnus atratus)

We studied patterns of parentage in 85 broods (332 cygnets) of black swans during three breeding seasons, using a set of eight polymorphic microsatellite markers. We detected both intraspecific brood parasitism (IBP; < 5% of cygnets per year) and extra-pair paternity (EPP). In these years, 10-17% (mean = 15.1%) of cygnets resulted from EPP, and 27-40% (mean 37.6%) of broods contained at least one extra-pair cygnet. Compared with levels of EPP in closely related species with similar life histories, these values are unexpectedly high. EPP in black swans appears unrelated to ecological factors (breeding density and synchrony) or genetic factors (genetic similarity between pair members or genetic quality of the offspring). We found no evidence that a mutual sexual feather ornament known to play a role in social mate choice in black swans (curled wing feathers) is involved in extra-pair mate choice. EPP does not lead to greater variance in reproductive success in males, relative to females in this species. We therefore suggest that EPP does not result in differential sexual selection on males and females, explaining why they are ornamented to the same degree.     

Avoid nest predation when predation rates are low,and other lessons: testing the tropical–temperate nest predation paradigm

Nest predation is the most important cause of nest failure in most birds and latitudinal differences in nest predation rates and life histories suggest that nest predation has been influential in life history evolution. All else equal, natural selection should favor reduction of nest predation, yet evidence is equivocal. We used Monte Carlo simulations to examine the combined effects of variation in nest predation rates, breeding season length and renesting intervals on the annual number of young fledged. Simulations suggest that selection most strongly favors a reduction in nest predation when breeding seasons are short and predation rates are low (temperate characteristics). Conversely, selection favors shorter renesting intervals when breeding seasons are long and nest predation rates are high (tropical characteristics). Reducing already low rates provides a proportionately greater increase in annual nesting success than does the same reduction when nest predation rates are higher. In some tropical species, individuals increase reproductive success not by avoiding predation in subsequent nesting attempts, which is largely beyond their control, but rather by reducing renesting intervals. We suggest that the emphasis on nest predation avoidance has biased our perspectives for alternative hypotheses of how birds should respond to nest predation and the consequences of those alternatives for life history theory. Similarly to the need to control for phylogenetics in examining life history strategies, future studies must also control for differences in breeding season lengths and renesting intervals to better understand the influence of nest predation on avian life histories.     

Interactive life‐history traits predict sensitivity of plants and animals to temporal autocorrelation

Temporal autocorrelation in demographic processes is an important aspect of population dynamics, but a comprehensive examination of its effects on different life‐history strategies is lacking. We use matrix population models from 454 plant and animal populations to simulate stochastic population growth rates (log λ_s) under different temporal autocorrelations in demographic rates , using simulated and observed covariation among rates. We then test for differences in sensitivities, or changes of log λ_s to changes in autocorrelation among two major axes of life‐history strategies, obtained from phylogenetically informed principal component analysis: the fast‐slow and reproductive‐strategy continua. Fast life histories exhibit highest sensitivities to simulated autocorrelation in demographic rates across reproductive strategies. Slow life histories are less sensitive to temporal autocorrelation, but their sensitivities increase among highly iteroparous species. We provide cross‐taxonomic evidence that changes in the autocorrelation of environmental variation may affect a wide range of species, depending on complex interactions of life‐history strategies.     

Extra-pair paternity and sexual dimorphism in birds

Differences in the strength of sexual selection between males and females can lead to sexual dimorphism. Extra-pair paternity (EPP) can increase the variance in male reproductive success and hence the opportunity for sexual selection. Previous research on birds suggests that EPP drives the evolution of dimorphism in plumage colour and in body size. Because EPP increases the intensity of sexual selection in males, it should lead to increased dimorphism in species with larger or more colourful males, but decreased dimorphism in species with larger or more colourful females. We explored the covariation between EPP and sexual dimorphism in wing length and plumage colouration in 401 bird species, while controlling for other, potentially confounding variables. Wing length dimorphism was associated positively with the frequency of EPP, but also with social polygamy, sex bias in parental behaviour and body size and negatively with migration distance. The frequency of EPP was the only predictor of plumage colour dimorphism. In support of our prediction, high EPP levels were associated with sexual dichromatism, positively in species in which males are more colourful and negatively in those in which females are more colourful. Contrary to our prediction, high EPP rates were associated with increased wing length dimorphism in species with both male- and female-biased dimorphism. The results support a role for EPP in the evolution of both size and plumage colour dimorphism. The two forms of dimorphism were weakly correlated and predicted by different reproductive, social and life-history traits, suggesting an independent evolution.     

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.     

Genetic variation and reproductive system among North American species of Nuttallanthus (Plantaginaceae)

We examined the effect of reproductive and life history strategies on the amount and partitioning of genetic variation in three annual species of Nuttallanthus. The North American species N. canadensis, N. floridanus, and N. texanus have regional to widespread ranges that overlap in the southeastern USA, are characterized by homogeneous populations and high fecundity, and possess showy, fragrant flowers seemingly adapted for insect pollination and outbreeding. Field and greenhouse studies on plants from 25 populations indicated that reproductive strategies were similar among species and showed predominant self-fertilization via cleistogamy and self-pollination prior to anthesis in chasmogamous flowers. Species were reproductively isolated and demonstrated complete cross-incompatibility after experimental crosses and no evidence for hybridization in mixed populations. Genetic variation was assessed using starch gel electrophoresis to resolve 15 isozyme loci in 50 populations. Conspecific genetic identity (I) values were high (0.819-0.936), but interspecific comparisons indicated many qualitative allelic differences and correspondingly low I values (0.516-0.623). Low levels of polymorphism and observed heterozygosity within populations and the disproportionate amount of gene diversity distributed among populations were concordant with reproductive data. The pattern of genetic differentiation was most similar to that observed in species with a predominantly inbreeding mating system.     

Genetic diversity does not explain variation in extra‐pair paternity in multiple populations of a songbird

Many songbirds are socially monogamous but genetically polyandrous, mating with individuals outside their pair bonds. Extra‐pair paternity (EPP) varies within and across species, but reasons for this variation remain unclear. One possible source of variation is population genetic diversity, which has been shown in interspecific meta‐analyses to correlate with EPP but which has limited support from intraspecific tests. Using eight populations of the genetically polyandrous red‐winged blackbird (Agelaius phoeniceus), including an island population, we investigated whether population‐level differences in genetic diversity led to differences in EPP. We first measured genetic diversity over 10 microsatellite loci and found, as predicted, low genetic diversity in the island population. Additional structure analyses with multilocus genotypes and mtDNA showed the island population to be distinct from the continental populations. However, the island population's EPP rate fell in the middle of the continental populations' distribution, whereas the continental populations themselves showed significant variation in EPP. This result suggests that genetic diversity by itself is not a predictor of EPP rate. We discuss reasons for the departure from previous results, including hypotheses for EPP that do not solely implicate female‐driven behaviour.     

Reproductive investment and somatic growth rates in longear sunfish

Synopsis Allocation of energy to current reproduction at the expense of other functions, such as growth, can limit future reproductive potential. This cost of reproduction is a central concept of life history theory but has been difficult to verify in comparative field studies. Three levels of comparison of growth rates and reproductive investments were evaluated within and among populations of longear sunfish,Lepomis megalotis. All three demonstrated high levels of reproductive investment associated with reduced somatic growth. Within populations of central longear sunfish there are precociously mature sneaker makes and later maturing parental makes; sneakers have greater gonadosomatic index (GSI) values and slower somatic growth rates than parental makes. Between subspecies of longear sunfish grown under common conditions, there are differences in age at maturity and in the level of physiological reproductive investment that are associated with distinct differences in growth rates. Between populations of central longear sunfish inhabiting different sites, there are differences in the level of reproductive investment that are also associated with differences in somatic growth. Each comparison produced evidence that trade-offs occur between these life history traits, supporting the hypothesis that there is a cost of reproduction among male sunfish and suggesting that differences in strategies of reproductive investment contribute to variation in somatic growth.     

Interspecific variation in plumage colour among birds: species recognition or light environment?

Abstract The traditional explanation for interspecific plumage colour variation in birds is that colour differences between species are adaptations to minimize the risk of hybridization. Under this explanation, colour differences between closely related species of birds represent reproductive character displacement. An alternative explanation is that interspecific variation in plumage colour is an adaptive response to variation in light environments across habitats. Under this explanation, differences in colour between closely related species are a product of selection on signal efficiency. We use a comparative approach to examine these two hypotheses, testing the effects of sympatry and habitat use, respectively, on divergence in male plumage colour. Contrary to the prediction of the Species Isolation Hypothesis, we find no evidence that sympatric pairs of species are consistently more divergent in coloration than are allopatric pairs of species. However, in agreement with the Light Environment Hypothesis, we find significant associations between plumage coloration and habitat use. All of these results remain qualitatively unchanged irrespective of the statistical methodology used to compare reflectance spectra, the body regions used in the analyses, or the exclusion of areas of plumage not used in sexual displays. Our results suggest that, in general, interspecific variation in plumage colour among birds is more strongly influenced by the signalling environment than by the risk of hybridization.     

Cross‐amplification of polymorphic microsatellites reveals extra‐pair paternity and brood parasitism in Sturnus vulgaris

We tested the cross‐amplification of 37 microsatellites in a population of starlings (Sturnus vulgaris). Twenty‐three of them amplified and five exhibited a large number of alleles per locus and high heterozygosity (on average: 14.6 alleles/locus and H_E = 0.704). We assessed the occurrence of extra‐pair paternity (EPP) and intraspecific brood parasitism (IBP) in this population. The EPP rate was 16% to 18% offspring from 43% to 45% of nests. IBP was very variable between two successive years (14% to 27% chicks from 25% to 64% of clutches). These five polymorphic markers will be of potential use in studies of genetic diversity, population structure and reproductive strategy of this species.     

Ecological significance of leaf life span among Central European grass species

Interspecific variation in leaf life span reflects the variation in nutrient conservation ability among different plant species and is considered to be associated with nutrient availability in the characteristic habitat. As defoliation interferes with nutrient conservation by the long-lived leaves, we hypothesized that disturbance rate is another important environmental factor working as a selective force on interspecific variation in leaf life span. In order to investigate this, we measured leaf life span of 32 grass species in mature garden-grown individuals. Variation in leaf life span was compared to measured leaf traits, to available data on species occurrence along gradients of nutrient availability and disturbance, and to published relative growth rates of the species. Leaf life span was associated positively with leaf tissue mass density and negatively with specific leaf area. Leaf life span correlated negatively with the disturbance rate in the characteristic habitat of a species, but not with nutrient availability. The latter relationship did not come about due to the long leaf life spans of species from nutrient-rich habitats with a relatively low disturbance rate, and to some extent also due to the short leaf life spans of annual species from relatively nutrient-poor sites. We conclude that although leaf longevity is an important means of reducing nutrient losses, this is a selective advantage only if the plant is not subjected to frequent defoliation. The frequently postulated association between leaf life span of a species and nutrient availability in its characteristic habitat may occur among species of habitats with positively correlated nutrient availability and disturbance rate. Leaf life span is negatively associated with seedling RGR, but there may be deviations in this relationship due to species with contrasting characteristics at seedling stage and at maturity.     

Evolution of passerine incubation behavior: influence of food, temperature, and nest predation

Abstract.— Incubation behavior is one component of reproductive effort and thus influences the evolution of life-history strategies. We examined the relative importance of body mass, frequency of mate feeding, food, nest predation, and ambient temperature to explain interspecific variation in incubation behavior (nest attentiveness, on- and off-bout durations, and nest trips per hour) using comparative analyses for North American passerines in which only females incubate. Body mass and frequency of mate feeding explained little variation in incubation behavior. We were also unable to detect any influence of food; diet and foraging strategy explained little interspecific variation in incubation behavior. However, the typical temperature encountered during reproduction explained significant variation in incubation behavior: Species breeding in colder environments take shorter bouts off the nest, which prevents eggs from cooling to temperatures below the physiological zero temperature. These species must compensate for shorter off-bouts by taking more of them (thus shorter on-bouts) to obtain needed energy for incubation. Nest predation also explains significant variation in incubation behavior among passerines: Species that endure high nest predation have evolved an incubation strategy (long on- and off-bouts) that minimizes activity that could attract predators. Nest substrate explained additional variation in incubation behavior (cavity-nesting birds have shorter on-bouts and make more frequent nest trips), presumably because nest predation and/or temperature varies among nest substrates. Thus, nest predation can influence reproductive effort in a way previously not demonstrated–by placing a constraint on parental activity at the nest. Incubating birds face an ecological cost associated with reproductive effort (predation of entire brood) that should be considered in future attempts to explain avian life-history evolution.     

Variability in swimming performance and underlying physiology in rainbow trout (Oncorhynchus mykiss) and brown trout (Salmo trutta)

We investigated intra- and interspecific variation in swimming performance and related physiological parameters in two members of the salmonid family. For our comparisons, we sourced juvenile brown trout (Salmo trutta) and rainbow trout (Oncorhynchus mykiss) from one hatchery and a second strain of rainbow trout from another. The hatcheries maintain genetic stocks obtained several decades ago from very different environments. We tested competing hypotheses: that there would be greater interspecific (across species) variation or that there would be greater intraspecific (within species) variation, owing to regional adaptations. To test these hypotheses, individual and small schools of five fish were taken to fatigue using the critical swimming speed test (U(crit)), and three post-exercise physiological metrics, packed red cell volume (hematocrit), citrate synthase and lactate dehydrogenase activity, were assessed. The majority of the results in swimming performance and hematocrit support that intraspecific variation was greater than interspecific variation, i.e. the location had a stronger effect than did genus. Variation in lactate dehydrogenase activity supported neither intra- nor interspecific variation as determining factors. In sum, our findings suggest that the performance of different species of salmonids from the same locale can be more similar than those of the same species from different areas.