No evidence for extra-pair paternity in the western gull

The genetic mating system of western gulls Larus occidentalis breeding on Southeast Farallon Island, California, was determined using multilocus DNA fingerprints of 33 chicks from 22 broods. No extra-pair paternity (EPP) was found, despite extra-pair copulations (EPCs) occurring. This suggests that paternity guards are effective, and that females gain few genetic benefits from EPCs. The EPP in western gulls concurs with that of other seabirds, reinforcing the idea that seabirds generally have a monogamous genetic mating system.     

No evidence of extra-pair paternity in the little auk Alle alle

Extra-pair paternity is typically rather infrequent in seabirds, as in most other long-lived and socially monogamous birds. Here we report the first paternity study of the little auk Alle alle , a high arctic seabird which raises only a single chick per year. Parentage was determined using three highly polymorphic microsatellite markers. We found that all 26 chicks in our sample were true genetic offspring of their social parents, with an upper 95% confidence limit of 10.88% for the frequency of extra-pair paternity. This level of extra-pair paternity is not significantly different from frequencies reported from the closely related common Uria aalge and Brünnich's guillemots U. lomvia .     

Genetic evidence for high propagule pressure and long‐distance dispersal in monk parakeet (Myiopsitta monachus) invasive populations

The monk parakeet (Myiopsitta monachus) is a successful invasive species that does not exhibit life history traits typically associated with colonizing species (e.g., high reproductive rate or long‐distance dispersal capacity). To investigate this apparent paradox, we examined individual and population genetic patterns of microsatellite loci at one native and two invasive sites. More specifically, we aimed at evaluating the role of propagule pressure, sexual monogamy and long‐distance dispersal in monk parakeet invasion success. Our results indicate little loss of genetic variation at invasive sites relative to the native site. We also found strong evidence for sexual monogamy from patterns of relatedness within sites, and no definite cases of extra‐pair paternity in either the native site sample or the examined invasive site. Taken together, these patterns directly and indirectly suggest that high propagule pressure has contributed to monk parakeet invasion success. In addition, we found evidence for frequent long‐distance dispersal at an invasive site (～100 km) that sharply contrasted with previous estimates of smaller dispersal distance made in the native range (～2 km), suggesting long‐range dispersal also contributes to the species’ spread within the United States. Overall, these results add to a growing body of literature pointing to the important role of propagule pressure in determining, and thus predicting, invasion success, especially for species whose life history traits are not typically associated with invasiveness.     

The importance of genetic evidence for identifying intra-specific brood parasitism

Intra-specific brood parasitism is widespread among birds. When genetic evidence is not available, criteria like super-normal clutch size, high within-clutch variance in egg morphology, and shorter than normal laying intervals have been used to identify parasitized broods. Here we report genetically determined parentage of a clutch of super-normal size in a species with fixed clutch size, the northern lapwing Vanellus vanellus . Results from multi-locus DNA fingerprinting revealed no loss of parentage in this brood. Furthermore, no loss of parentage was found in another clutch with a high variance in egg size and colouration. These findings add to previous evidence that reliable classification of brood parasitism requires molecular analysis.     

Ecological constraints on extra-pair paternity in the bluethroat

The mating system of the bluethroat (Luscinia s. svecica) involves a high level of sperm competition, and consequently a high frequency of extra-pair paternity (EPP). There is considerable variation in the frequency of EPP. Over the course of ten study years in a population in Norway, the frequency has fluctuated between 7% and 33% of young, and 8% and 76% of broods. In this paper, we address the issue of whether ecological factors can explain some of the variation in EPP between years and broods. Factors include breeding density, breeding synchrony and two meteorological variables (ambient temperature and precipitation) during the period of peak female fertility. There were no significant relationships between annual averages of the four variables and the annual level of EPP, but the statistical power of these tests was low, due to a restricted sample (n=10 years). Focussing on individual broods, none of the factors had significant effects when including all broods in the analyses (including those with zero EPP). When limiting the analyses to broods with one or more extra-pair offspring, morning temperature had a significant effect on the frequency of EPP. When the temperature was low during the peak of the fertile period, EPP occurred less frequently. Male extra-pair behaviour was not generally constrained by synchrony between the fertile periods of the social mate and the extra-pair mate, but for a subgroup of males ("unattractive males") there appeared to be a trade-off between mate guarding and pursuing extra-pair copulations. Our results indicate that ecological conditions have no influence on whether or not a brood will contain extra-pair offspring, but when EPP does occur, its frequency is influenced by air temperature. Possibly, there is a causal relationship between weather conditions and the frequency of extra-pair encounters in this species. The breeding season is initiated at a phenologically early stage, when large parts of the territories are still covered with snow, and cold weather conditions may force the birds to devote most of their time to maintenance and thereby constrain sexual activities.     

Assessing the potential range expansion of the exotic monk parakeet in Spain

In this study we determine favourable areas for the monk parakeet, Myiopsitta monachus , in peninsular Spain to account for its current distribution and predict its future course according to its potential range. We applied a favourability function based on generalized linear models using the presence/absence of breeding colonies of the species and the values of a set of variables on the 5167 UTM 10 × 10 km squares comprising the study area. We calculated the factor of distribution change in presences predicted by the model, and grouped the variables into explanatory factors performing a variation partitioning to assess the explanatory power of each factor. Our model included six predictors to explain the presence and absence of the species. These predictors were grouped into three factors: human activity, climate, and topography. Purely human influences accounted for 63.8% of the variation of the final model, while topographical variables explained 15.2% and climate only 5.7%. We obtained a high distribution change factor in which the presences of the species were predicted to increase between two- and sevenfold. Taking into account highly favourable squares, we conclude that the species is still absent in more than 72% of potential settlement areas, and thus we expect a continuous increase in the distribution of the species. Human activity is the main force moulding the distribution of the species, and lies behind its fast expansion, which is not only active, but is also passive via releases and escapes. We identified the areas of likely future expansion of the exotic monk parakeet in Spain. The pest status of the species in its native range, together to its distribution trend, should be taken into account by wildlife agencies to consider options for management.     

Egg patterning is not a reliable indicator of intraspecific brood parasitism in the blue tit Cyanistes caeruleus

A recent study reported the incidence of intraspecific brood parasitism (IBP) in the blue tit, based on differences in pigmentation between eggs within clutches and the appearance of multiple eggs within a nest in a single day. Here we present data from another population of blue tits Cyanistes caeruleus in which we show that such evidence is not necessarily indicative of IBP. As in many other studies of birds, we found that the patterning and degree of pigmentation on eggs is individually highly characteristic of a female. As in the previous study of blue tits, a minority of clutches (about 3% of 904 clutches) contained one, two or three eggs that were distinctly different from the other eggs in the clutch, either in size or patterning, but not both. Molecular genetic analyses of parentage in a sample of these clutches with suspected IBP demonstrated unequivocally that all eggs within each clutch had in fact been laid by a single female, in all cases the social parent. We also found no evidence of IBP from paternity analyses of a sample of 3,529 offspring from 427 broods in this population, adding to previous evidence of the absence of IBP in this species. These findings cast doubt on the utility of purely observational approaches to provide evidence of intraspecific brood parasitism; our study suggests that egg morphology and records of laying patterns are inadequate to enable the identification of IBP.     

Hindlimb myology of the monk parakeet (Aves,Psittaciformes)

We studied the hindlimb myology of the monk parakeet (Myiopsitta monachus). Like all parrots, it has zygodactyl feet enabling perching, climbing, hanging, moving easily among trees, and handling food. Muscles were described and weighed, and physiological cross‐sectional area (PCSA) of four flexors and one extensor was calculated. In comparison to other muscles, the M. tibialis cranialis and the M. fibularis brevis show increased development and high PCSA values, and therefore, large potential force production. Also, a large proportion of muscle mass was involved in flexing the digits. We hypothesize that these muscle traits are associated with the arboreal locomotion and food manipulation habits. In the monk parakeet, the M. extensor digitorum longus sends a branch to the hallux, and the connection between the M. flexor digitorum longus and the M. flexor hallucis longus is type I (Gadow's classification). We reaffirm the presence of the M. ambiens as a plesiomorphic condition that disappears in most members of the order. Among Psittaciformes, the M. fibularis brevis is stronger and the M. fibularis weaker in arboreal species than in basal terrestrial ones (e.g., Strigops). J. Morphol. 275:732–744, 2014. © 2014 Wiley Periodicals, Inc.     

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.     

Relatedness and the evolution of conspecific brood parasitism

In conspecific brood parasitism (CBP), a parasitic female takes advantage of the parental care performed by a host female by laying eggs in the nest of the host. The host female raises the offspring of the parasitic female as well as her own. In species where local females are related, direct costs for the host might be more than compensated for by gains in inclusive fitness through increased reproduction of a related parasite, but the role of relatedness in CBP is debated. This inclusive-fitness model of parasitism, structured as a game between host and parasite, suggests that both females can gain inclusive fitness and that host-parasite relatedness can therefore facilitate the evolution of CBP. Crucial assumptions are that there is kin discrimination and a potential for host resistance to parasitism by unrelated females but close relatives are accepted. The cost of parasitism in terms of reduced clutch size or offspring survival for the host must not be large; otherwise, parasitism will reduce her inclusive fitness. Therefore, if these costs are high, it does not benefit a host to accept a parasite, even if the parasite is closely related. The secondary female may still have higher fitness from parasitism, but if the costs are high, she should parasitize an unrelated host, not a relative. This requires that the reduction in parasite success that a host can cause by resistance is not too large; otherwise, it will be better for the secondary female to parasitize an accepting related host or to nest solitarily. For these reasons, host-parasite relatedness is most likely to occur in animals where costs of being parasitized are low and host resistance can markedly reduce the success of an unrelated parasite. When costs are higher, parasitism of unrelated hosts may be better, and if host resistance strongly reduces parasite success, solitary breeding is preferable. In some cases, CBP is directly advantageous for the host, and it may sometimes evolve in close connection with cooperative breeding, which is also considered in the model. Some but not all empirical results support these ideas, and more detailed studies of behavior, relatedness, and reproduction of host and parasite are needed for critical tests.     

Conspecific brood parasitism and population dynamics

Conspecific brood parasitism (CBP), defined as parasitic laying of eggs in a conspecific nest without providing parental care, occurs in insects, fishes, amphibians, and many birds. Numerous factors have been proposed to influence the evolution of CBP, including nest site limitation; effects of brood size, laying order, or parasitic status on offspring survival; randomness of parasitic egg distribution; adult life-history trade-offs; and variation in parental female quality or risk of nest predation. However, few theoretical studies consider multiple possible types of parasitism or the interplay between evolution of parasitism and population dynamics. We review existing theory of CBP and develop a synthetic modeling approach to ask how best-of-a-bad job parasitism, separate-strategies parasitism (in which females either nest or parasitize), and joint-strategies parasitism (in which females can both nest and parasitize) differ in their evolutionary conditions and impacts on population dynamics using an adaptive dynamics framework including multivariate traits. CBP can either stabilize or destabilize population dynamics in different scenarios, and the role of comparable parameters on evolutionarily stable strategy parasitism rate, equilibrium population size, and population stability can differ for the different modes of parasitism.     

The evolution of brood parasitism: the role of facultative parasitism

The hypothesis that facultative brood parasitism may serve asan intermediate step in the evolutionary transition from purelyparental reproduction to obligate parasitism was investigated.The population dynamics of a host-parasite complex were computer-simulatedin a model that incorporated different intensities of parasitismand host defense and considered a simplified semelparous birdspecies living in a homogeneous habitat The individuals usetwo different breeding strategies: provide parental care orparasitize the nest of those providing parental care. Underobligate parasitism, the parasites appeared unsuccessful, drovethe host population to extinction, or coexisted with the hostin stable or oscillating proportions. The behavior of the systemdepended on both the effectiveness of the parasite and the defenseof the host. Under facultative parasitism (making the best ofa bad job), the parasites reduced host numbers but did not reducethe population size below the number of breeding sites. Thus,facultative parasitism provides a better opportunity for thedevelopment of defense in the host. The population of a hostthat shows a certain level of defense can be more successfullyinvaded by obligate parasites so that stable coexistence ofhosts and parasites is possible.     

Female choice and extra-pair paternity in a traditional human population

Seeking out extra-pair paternity (EPP) is a viable reproductive strategy for females in many pair-bonded species. Across human societies, women commonly engage in extra-marital affairs, suggesting this strategy may also be an important part of women's reproductive decision-making. Here, I show that among the Himba 17 per cent of all recorded marital births are attributed by women to EPP, and EPP is associated with significant increases in women's reproductive success. In contrast, there are no cases of EPP among children born into 'love match' marriages. This rate of EPP is higher than has been recorded in any other small-scale society. These results illustrate the importance of seeking EPP as a mechanism of female choice in humans, while simultaneously showing it to be highly variable and context-dependent.     

Interspecific brood parasitism in galliform birds

Mode of development in birds helps determine the form of brood parasitism a species exhibits. Most knowledge of precocial brood parasites comes from a single avian family, the waterfowl (Anatidae: Anseriformes). Here we review cases of interspecific brood parasitism (IBP) in a second group of precocial birds, the order Galliformes. IBP is uncommon but taxonomically widespread, occurring in at least 11 species and in four of five galliform families. By far the most common brood parasite is the Ring-necked Pheasant Phasianus colchicus . Hosts were generally other ground-nesting precocial species. It is unclear whether the absence of IBP in the Cracidae (Guans, Curassows, and Chachalacas) is due to the paucity of research on tropical gamebirds or because tropical birds such as the Cracidae may be less likely to practise IBP. Galliform birds mirror the trend found in ducks in which virtually all species that parasitize heterospecifics are also conspecific brood parasites (CBP). This association supports the hypothesis that IBP as an adaptive tactic or strategy may evolve from CBP. Alternatively, or additionally, egg-dumping may represent reproductive error on the part of females, such that concordance between CBP and IBP could be a byproduct of having sufficient knowledge of breeding biology only for a subset of galliform species.     

Social structure influences extra-pair paternity in socially monogamous mammals

Using the genetic estimates of paternity available for 22 species of socially monogamous mammals, we investigated the impact of the social structure and of the type of pair bonding on the interspecific variations of extra-pair paternity rates. To this purpose, we classified species in three categories of social structure—solitary, pair or family-living species—and in two categories of pair bonding—intermittent or continuous. We show that interspecific variations of extra-pair paternity rates are better explained by the social structure than by the type of pair bonding. Species with intermittent and continuous pair bonding present similar rates of extra-pair paternity, while solitary and family-living species present higher extra-pair paternity rates than pair-living species. This can be explained by both higher male–male competition and higher female mate choice opportunities in solitary and family-living species than in pair-living species.     

Modelling the arms race in avian brood parasitism

In brood parasitism, interactions between a parasite and its host lead to a co-evolutionary process called an arms race, in which evolutionary progress on one side provokes a further response on the other side. The host evolves defensive means to reduce the impact of parasitism, while the parasite evolves means to counter the host's defence. To gain insights into the co-evolutionary process of the arms race, a model is developed and analysed, in which the host's defence and the parasite's counterdefence are assumed to be genetically determined. First, the effect of parasite counterdefence on host defence is analysed. I show that parasite counterdefence can critically affect the establishment of host defence, giving rise to three situations in the equilibrium state: The host shows (1) no defence, (2) an intermediate level of defence or (3) perfect defence. Based on these results, the evolution of parasite counterdefence is considered in connection with host defence. It is suggested that the parasite can evolve counterdefence to a certain degree, but once it has established counterdefence beyond this, the host gives up its defence against parasitism provided the defence entails some cost to perform. Dynamic aspects of selection pressure are crucial for these results. Based on these results, I propose a hypothetical evolutionary sequence in the arms race, along which interactions between the host and parasite proceed.     

Diet specialization and brood parasitism in cuckoo species

1. Brood parasitism is a breeding strategy adopted by many species of cuckoos across the world. This breeding strategy influences the evolution of life histories of brood parasite species.
2. In this study, we tested whether the degree on diet specialization is related to the breeding strategy in cuckoo species, by comparing brood parasite and nonparasite species. We measured the gradient of diet specialization of cuckoos, by calculating the Gini coefficient, an index of inequality, on the multiple traits describing the diet of species. The Gini coefficient is a measure of statistical dispersion on a scale between 0 and 1, reflecting a gradient from low to high specialization, respectively. First, we tested the strength of the phylogenetic signal of diet specialization index among cuckoo species worldwide. Then, we ran phylogenetic generalized least square (PGLS) models to compare diet specialization, distribution range, and body mass of parasitic and nonparasitic cuckoo species, considering the phylogenetic signal of data.
3. After adjusting for the phylogenetic signal of the data and considering both, species distribution range and species body mass, brood parasitic cuckoos were characterized by higher diet specialization than nonbrood parasitic species. Brood parasitic species were also characterized by a larger breeding distribution range than nonparasitic species.
4. The findings of this study provide an additional understanding of the cuckoos’ ecology, relating diet and breeding strategies, information that could be important in conservation ecology.