Rozprawy doktorskie na temat „Extra-pair paternity”

Thesis (Ph. D.) -- University of Maryland, College Park, 2006.
Thesis research directed by: Behavior, Ecology, Evolution and Systematics. Title from t.p. of PDF. Includes bibliographical references. Published by UMI Dissertation Services, Ann Arbor, Mich. Also available in paper.

Mate competition is a prominent component of sexual selection theory. Typically, males attempt to mate with the most females possible and females attempt to mate with the highest quality males possible. In the polygynous female-defense mating system of Anolis carolinensis, males compete directly for females through territorial behavior. Inter-male competition is intense due to an average polygyny ratio of 1 male to 3 females despite a 1:1 adult sex ratio. Through high levels of territorial behavior (e.g., 100 displays/h, 27 m patrol distances/h, 70% of day in defense-related activities), males attempt to exclude other males from resident females who, in turn, both store sperm and ovulate a single-egg clutch at weekly intervals over a 4-month breeding season. Paternity of hatchlings in 16 naturally occurring breeding groups was analyzed to determine the extent to which the territorial resident male was able to prevent other males from fathering offspring of his resident females. Lizards residing in or neighboring a resident maleâ s territory were collected and RAPD-PCR was used to determine the paternity of hatchlings. Of the 48 hatchlings from 26 females, resident territorial males fathered 52% of hatchlings; 15% were fathered by a male whose territory bordered that of the resident male and 21% were fathered by a smaller male living covertly within the resident maleâ s territory. Paternity for the remaining 12% of hatchlings belonged to an unsampled male. Given that females mated with multiple males, laboratory-based controlled matings were conducted where females were sequentially paired with two males and RAPD-PCR was used to analyze which of the two males fathered the subsequent hatchlings to determine the mechanism of sperm precedence. Regardless of mating order, only one male of the pair fertilized the eggs. Male A. carolinensis have reproductive strategies present in addition to defending resident females and female A. carolinensis have options in addition to simply mating with the resident male. While sperm precedence is present in this species, it is not based on mating order, but may involve both the number of sperm deposited in the femaleâ s tract as well as the quality of those sperm.
Ph. D.

Apesar de a monogamia social ser dominante entre as aves, análises genéticas revelaram relações de parentesco inesperadas, evidenciando diferentes estratégias de reprodução, como a paternidade extra-par e o parasitismo de ninho. Espécies de passeriformes estão entre as mais promíscuas, com altas taxas de paternidade extra-par, enquanto em aves marinhas esse comportamento demonstrou ser menos frequente. Pinguins (Família Spheniscidae) compõem um grupo de 18 espécies de aves marinhas pelágicas e que tem em comum a filopatria, fidelidade a um parceiro e intenso cuidado biparental. Portanto, espera-se que apresentem um comportamento estritamente monogâmico e taxas de paternidade extra-par insignificantes. Avaliamos pela primeira vez o sistema reprodutivo dos Pinguins-de-Magalhães através de uma abordagem genética, buscando investigar a existência e frequência de paternidade extra-par e parasitismo de ninho. O parentesco de 88 filhotes de 44 ninhos de uma colônia na Ilha Quiroga (Argentina) foi determinado com base em análises de 9 marcadores microssatélites. Encontramos baixas taxas de parasitismo de ninho (6%), mas altas taxas de paternidade extra-par (31% e 48% dos ninhos com pelo menos 1 filhote extra-par). Entre os dois anos coletados, encontramos uma pequena diferença na incidência de infidelidade (29% em 2010; 32% em 2011), mas não houve relação com as condições climáticas do período de reprodução da espécie. Além disso, apesar da alta taxa de filhotes extra-par, não encontramos diferença significativa na diversidade genética e nem viés da razão sexual secundária. Acreditamos que a alta taxa de paternidade extra-par encontrada possa ter relação com o comportamento reprodutivo em colônia, a densidade populacional, o sincronismo reprodutivo, ou que parte da paternidade que não correspondeu aos pais sociais seja resultado de troca de parceiros antes da definição final dos casais em cada estação reprodutiva. Nosso estudo pode ajudar a melhor entender e caracterizar o sistema reprodutivo dos Pinguins-de-Magalhães e indica que a espécie é socialmente, mas não geneticamente monogâmica.
Despite the social monogamy being dominant among birds, genetic analysis revealed unexpected kinship relations, showing different reproductive strategies, such as extra-pair paternity and brood parasitism. Passerine species are among the most promiscuous, with high extra-pair paternity rates, while in seabirds this behavior is typically rather less frequent. Penguins (Spheniscidae Family) are a group of 18 species of pelagic seabirds that have in common philopatric behavior, faithfulness to one partner and intense biparental care. Therefore, they are expected to have a strictly monogamous behavior and insignificant rates of extra-pair paternity. For the first time, we evaluated the reproductive system of Magellanic Penguins (Spheniscus magellanicus) through genetic analysis in order to investigate the existence and frequency of extra-pair paternity and brood parasitism. The kinship of 88 offspring of 44 nests from a colony on Quiroga Island (Argentina) was determined based on the analyses of 9 microsatellite markers. We found low rates of brood parasitism (6%), but high extra-pair paternity rates (31% and 48% of nests with at least one extra-pair offspring). Between the two years sampled, we found a small difference in the incidence of infidelity (29% in 2010; 32% in 2011), but no connection with the climatic conditions of each breeding season. In addition, despite the high rate of extra-pair offspring, we found no significant difference in the genetic diversity and no bias in the secondary sex ratio. We believe that the high rate of extra-pair paternity found in our study may be a result of their reproductive behavior of nesting in colonies, breeding synchrony, density, or that part of the mismatching paternity is due mate switching. Our study may help to better understand and characterize the reproductive system of Magellanic penguins and indicates that this species is socially but not sexually monogamous.

The Australian magpie (Gymnorhina tibicen) is a large, sedentary, omnivorous passerine. In some populations, individuals live in groups, and some of these groups breed cooperatively. The white-backed magpie (G. t. tyrannica) from the south-eastern corner of the continent, has had relatively little study, and few details are known of its mating system, social structure, and method of parental care. I conducted an observational study on a population of white-backed magpies, recording details of their demography, dispersal, breeding system, and parental care. In conjunction, I conducted a genetic analysis of the population, to determine if the genetic mating system matched the observed social system, to detect instances of extra-group mating, and to sex juvenile birds. Extra-pair paternity (EPP) is a common feature of the mating systems of many birds. The rate of EPP may vary between species, races and populations. I made a comparison of extra-group paternity (EGP) rates between two races of the Australian magpie, to determine if similar mating systems were being employed. The two populations had similar social structure, but differed in group size and dispersal. I predicted that dispersal differences would have a profound effect on the rate of EGP between the populations, as the population with the lower rate of dispersal and higher chance of breeding with a close relative would engage in EGPs more frequently. Eight microsatellite loci were used to determine parentage in the white-backed Australian magpie. The rate of EGP was found to be 44%. Dispersal rates were estimated from observational data. Over half of the juvenile magpie cohort from the previous breeding season leave the territorial group. These results contrast sharply with the results found by other researchers in a population of western Australian magpies (G t. dorsalis). In this population, 82% EGP is recorded and dispersal of juveniles is close to nil. The results indicate that dispersal rate is a potentially important predictor of rates of extra-group fertilisations between populations of this species, and suggest that females maximise their reproductive output by avoiding breeding with close kin. The reproductive success of a male bird is often correlated with measurable traits that predict his intrinsic quality. Females are thought to select mates based on their quality to gain their 'good genes'. Male Australian magpies of the white-backed race were trapped in two breeding seasons. Measurements were taken of morphometric and other characteristics in order to discover whether particular traits of males were associated with: a) number of fledglings produced in the territory per season; b) percentage of offspring sired in the territory; and c) whether females select males for their 'good genes'. The only variable that was correlated with number of territorial offspring was feather lice load. Males with high numbers of lice were less likely to produce territorial fledglings in one season and across both seasons. Males of inferior quality may be subject to increased conspecific territorial intrusions, leading to more time spent on defence, more failed breeding attempts, less time allocated to grooming and thus high parasite loads. Males that produced many territorial fledglings were more likely to gain genetic paternity of at least some of them, although again this was significant for only one season. Also, across both seasons, a high number of females in the group was correlated with increased paternity within the group. The general lack of correlation between the variables and level of genetic paternity may be due to females engaging in extra-group mating primarily to avoid breeding with a close relative rather than to choose a quality male. In this scenario, males would not have to be 'high quality', but merely genetically different to the female's social mate. Extra-group paternity (EGP) can affect paternal effort. It may also influence the helping effort of auxiliary birds in cooperatively breeding species. If helping is driven by kin selection, helpers should decline to provision unrelated young. Relatedness becomes difficult to assess however, when females mate outside the group. Alternative rewards may then become important in helper decisions. In my study population of Australian magpies, 38% of fledglings were sired by males outside the territorial group. In a second population (G. t. dorsalis), 82% of fledglings were sired by extra-group males. I observed within-group male and helper feeding effort over three breeding seasons in the first population and obtained data recorded over a single season in the second population. In both populations, males provisioned young regardless of relatedness, as did helpers. Males provisioned less than the nesting female on average. Paternal effort did not reduce with an increase in the rate of EGP between populations. In the population with intermediate levels of EGP, the white-backed magpies, I observed helpers in about half of the sampled territories that produced fledglings. Helpers did not increase the production of young. In the population with high levels of EGP, western magpies, I detected helping behaviour in proportionally more territories. It appears that Australian magpie helpers provide help in order to pay 'rent' and remain on the natal territory. I discuss these results in light of the differences between the two races of magpie and the major theories regarding male parenting decisions and helper activity. Finally, I examine the relatively high rates of EGP's in the Australian magpie from a phylogenetic perspective. Although inbreeding avoidance is strongly supported by this study as the major reason EGP is so common in magpie populations, there may be an element of phylogenetic inertia that maintains the frequency of this behavioural trait. I comment upon the use of single-population estimates of species EGP rates in comparative analyses, given the intraspecific variation discovered between Australian magpie populations. Future directions for the study of mate choice in the Australian magpie are outlined with a proposal to study variation at the major histocompatibility complex between mated pairs.
Thesis (PhD Doctorate)
Doctor of Philosophy (PhD)
Australian School of Environmental Studies
Full Text

The Australian magpie (Gymnorhina tibicen) is a large, sedentary, omnivorous passerine. In some populations, individuals live in groups, and some of these groups breed cooperatively. The white-backed magpie (G. t. tyrannica) from the south-eastern corner of the continent, has had relatively little study, and few details are known of its mating system, social structure, and method of parental care. I conducted an observational study on a population of white-backed magpies, recording details of their demography, dispersal, breeding system, and parental care. In conjunction, I conducted a genetic analysis of the population, to determine if the genetic mating system matched the observed social system, to detect instances of extra-group mating, and to sex juvenile birds. Extra-pair paternity (EPP) is a common feature of the mating systems of many birds. The rate of EPP may vary between species, races and populations. I made a comparison of extra-group paternity (EGP) rates between two races of the Australian magpie, to determine if similar mating systems were being employed. The two populations had similar social structure, but differed in group size and dispersal. I predicted that dispersal differences would have a profound effect on the rate of EGP between the populations, as the population with the lower rate of dispersal and higher chance of breeding with a close relative would engage in EGPs more frequently. Eight microsatellite loci were used to determine parentage in the white-backed Australian magpie. The rate of EGP was found to be 44%. Dispersal rates were estimated from observational data. Over half of the juvenile magpie cohort from the previous breeding season leave the territorial group. These results contrast sharply with the results found by other researchers in a population of western Australian magpies (G t. dorsalis). In this population, 82% EGP is recorded and dispersal of juveniles is close to nil. The results indicate that dispersal rate is a potentially important predictor of rates of extra-group fertilisations between populations of this species, and suggest that females maximise their reproductive output by avoiding breeding with close kin. The reproductive success of a male bird is often correlated with measurable traits that predict his intrinsic quality. Females are thought to select mates based on their quality to gain their 'good genes'. Male Australian magpies of the white-backed race were trapped in two breeding seasons. Measurements were taken of morphometric and other characteristics in order to discover whether particular traits of males were associated with: a) number of fledglings produced in the territory per season; b) percentage of offspring sired in the territory; and c) whether females select males for their 'good genes'. The only variable that was correlated with number of territorial offspring was feather lice load. Males with high numbers of lice were less likely to produce territorial fledglings in one season and across both seasons. Males of inferior quality may be subject to increased conspecific territorial intrusions, leading to more time spent on defence, more failed breeding attempts, less time allocated to grooming and thus high parasite loads. Males that produced many territorial fledglings were more likely to gain genetic paternity of at least some of them, although again this was significant for only one season. Also, across both seasons, a high number of females in the group was correlated with increased paternity within the group. The general lack of correlation between the variables and level of genetic paternity may be due to females engaging in extra-group mating primarily to avoid breeding with a close relative rather than to choose a quality male. In this scenario, males would not have to be 'high quality', but merely genetically different to the female's social mate. Extra-group paternity (EGP) can affect paternal effort. It may also influence the helping effort of auxiliary birds in cooperatively breeding species. If helping is driven by kin selection, helpers should decline to provision unrelated young. Relatedness becomes difficult to assess however, when females mate outside the group. Alternative rewards may then become important in helper decisions. In my study population of Australian magpies, 38% of fledglings were sired by males outside the territorial group. In a second population (G. t. dorsalis), 82% of fledglings were sired by extra-group males. I observed within-group male and helper feeding effort over three breeding seasons in the first population and obtained data recorded over a single season in the second population. In both populations, males provisioned young regardless of relatedness, as did helpers. Males provisioned less than the nesting female on average. Paternal effort did not reduce with an increase in the rate of EGP between populations. In the population with intermediate levels of EGP, the white-backed magpies, I observed helpers in about half of the sampled territories that produced fledglings. Helpers did not increase the production of young. In the population with high levels of EGP, western magpies, I detected helping behaviour in proportionally more territories. It appears that Australian magpie helpers provide help in order to pay 'rent' and remain on the natal territory. I discuss these results in light of the differences between the two races of magpie and the major theories regarding male parenting decisions and helper activity. Finally, I examine the relatively high rates of EGP's in the Australian magpie from a phylogenetic perspective. Although inbreeding avoidance is strongly supported by this study as the major reason EGP is so common in magpie populations, there may be an element of phylogenetic inertia that maintains the frequency of this behavioural trait. I comment upon the use of single-population estimates of species EGP rates in comparative analyses, given the intraspecific variation discovered between Australian magpie populations. Future directions for the study of mate choice in the Australian magpie are outlined with a proposal to study variation at the major histocompatibility complex between mated pairs.

Inbreeding occurs when relatives mate and have offspring. Inbreeding depression is hypothesized to have influenced the evolution of mating systems and behavioural mechanisms of inbreeding avoidance in the animal kingdom. Inbreeding in the wild is difficult to measure, as in order to build a pedigree allowing us to identify matings between relatives, the identity of as many as possible members of a population needs to be known. For a long time, the main source of knowledge about inbreeding depression was based on laboratory and agricultural studies, which did not reflect the array of environmental pressures wild populations have to cope with. In consequence, the deleterious consequences of inbreeding have often been underestimated. This is problematic because accurate estimates of the effect size of inbreeding depression are needed to study the strength of selection on inbreeding avoidance mechanisms, and are also of importance to conservation genetics. The aim of this thesis was to use pedigree data to infer the occurrence and effects of inbreeding using over forty years of breeding events of the great tit Parus major from Wytham Woods, Oxfordshire. The effects of inbreeding on fitness were investigated across a life-history continuum, and across environments. I found that close inbreeding (f=0.25) resulted in pronounced inbreeding depression, which acted independently on hatching success, fledging success, and recruitment success, and reduced the number of fledged grand-offspring by 55%. My results therefore suggest that estimates of fitness costs of inbreeding must focus on the entire life cycle. I also show that the variation in the strength of inbreeding depression varies across environments, particularly so the more the environmental variable considered is linked to fitness. These results emphasise the need of using relevant environmental contrasts when investigating inbreeding by environment interactions. I further asked whether individuals involved in matings with relatives differed relative to individuals mating with unrelated partners. I did not find any evidence for clear predictors of inbreeding, and I show that inbreeding depression in our population is entirely independent of any tendency for low quality parental genotypes, or phenotypes, to inbreed. Neither did I find any evidence for active inbreeding avoidance: great tits did not mate less often with kin than expected based on several scenarios of random mating, nor did I find increased rates of extra-pair paternity among birds breeding with relatives. In fact, I observed quite the contrary, as birds mating with kin exhibited a higher than average rate of close inbreeding relative to all scenarios of random mating investigated, showed lower rates of extra-pair paternity and divorce than birds mated to unrelated partners. I hypothesise that cases of occasional inbreeding in this population may result from mis-imprinting or a related process whereby some birds develop particularly strong bonds that are at odds with all predictions of avoiding inbreeding. Finally, I asked to what extent natal dispersal, a behaviour that is often hypothesized to play an important role in avoiding inbreeding, indeed reduces the likelihood of inbreeding. I found that male and female individuals breeding with a relative dispersed over several-fold shorter distances than those outbreeding. This led to a 3.4 fold increase (2.3-5, 95% CI) in the likelihood of close inbreeding relative to the population average when individuals dispersed less than 200m. This thesis demonstrates that inbreeding has deleterious effects on a wild population of birds, occurring throughout an individual’s life, and is of varying strength across environments. My findings strongly support the theory that natal dispersal should be considered as a mechanism of prime importance for inbreeding avoidance.

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Universidade Federal de Sao Carlos
Population genetic parameters and genetic relatedness estimates were carried out for Roseate Spoonbill (Platalea ajaja), Wood Stork (Mycteria americana) and Great Egret (Ardea alba egretta) reproductive colonies in Amapa, Pantanal and Rio Grande do Sul (RS), Brazil. Microsatellite genotypes were used to investigate kinship patterns between nestlings sampled inside the same nests, using a variety of analytical approaches. Unrelated nestling-pairs were observed in Roseate Spoonbill nests (6.12% of analyzed nests) and in Wood Stork nests (11.34%); half-siblings were present in Roseate Spoonbill nests as well (1.36%). Only full-siblings were detected inside Great Egret nests. Conspecific brood parasitism (CBP) and extra-pair paternity were proposed to account for the presence of unrelated nestmates and half-siblings, respectively, in Roseate Spoonbill and Wood Stork nests. Those results suggest the occurrence of a mating system different than genetic monogamy in natural populations of those waterbirds. Genetic relatedness was also investigated for adults and offspring, as well as for supposed siblings in Roseate Spoonbill families kept in three zoological facilities in the U.S. Paternity and maternity allocation analyses through maximum-likelihood revealed that errors were present in zoo‟s studbooks in relation to the familial records. We also identified mating between related individuals that were not detected previously by zookeepers. Population genetic parameters were also estimated and demographic processes were assessed for Great Egret reproductive colonies in the Pantanal and Rio Grande do Su, Brazil. Bayesian clustering analyses, assignment tests, analysis of molecular variance, F-statistics estimates, allelic frequency distribution and the G-W index revealed that: i) Pantanal reproductive colonies are genetically differentiated from Rio Grande do Sul colonies; ii) an IBD-like pattern alone cannot explain that differentiation; and iii) genetic signal of a reduction of population size was present for two colonies in the Pantanal and one in Rio Grande do Sul. Results were discussed considering a metapopulation dynamic and also considering that populations from both Brazilian regions represent distinct units and deserve to be treated separately when planning and carrying out conservation and management programs that aim to preserve the species‟ genetic diversity.
Estudos de genética de populações e de parentesco genético foram desenvolvidos em colhereiro (Platalea ajaja), cabeça-seca (Mycteria americana) e garça-branca-grande (Ardea alba egretta), de colônias reprodutivas do Amapá, Pantanal e Rio Grande do Sul (RS), Brasil. Genótipos em locos de microssatélites foram utilizados para se investigar os padrões de relacionamento entre ninhegos amostrados dentro dos mesmos ninhos com diferentes metodologias de análise. Pares de ninhegos não-relacionados foram encontrados nos ninhos de colhereiro (6,12% dos pares analisados) e de cabeça-seca (11,34%); meio-irmãos foram observados nos ninhos de colhereiro (1,36%). Em garça branca grande foi detectada apenas a presença de irmãos-completos dentro dos ninhos. Parasitismo de ninho intraespecífico e paternidade extra-par podem explicar a presença de ninhegos não-relacionados e meio-irmãos nos ninhos de colhereiro e cabeça-seca, o que indica a presença de um sistema de acasalamento diferente da monogamia genética nas populações naturais dessas espécies. Relações de parentesco entre adultos e filhotes e entre supostos irmãos foram determinadas em famílias de colhereiro de três zoológicos dos EUA. Análises de atribuição de maternidade e paternidade por máxima verossimilhança revelaram erros nos registros dos zoológicos quanto às relações progenitor-progênie e identificaram acasalamentos entre indivíduos aparentados que não tinham sido registrados. Parâmetros genético-populacionais e processos demográficos foram investigados em populações de garça-branca-grande do Pantanal e do Rio Grande do Sul. Análises Bayesianas, testes de alocação de indivíduos, análises de variância molecular, estimativa de estatísticas F, exame da distribuição das freqüências alélicas e cálculo do índice de G-W permitiram identificar que: i) há diferenciação genética significativa entre colônias reprodutivas do Pantanal e do Rio Grande do Sul; ii) o padrão de isolamento pela distância não explica essa diferenciação; e iii) duas populações no Pantanal e uma população no RS apresentaram sinais genéticos de redução demográfica. Os resultados foram discutidos considerando que as populações de garça-branca-grande localizadas no Pantanal e no Rio Grande do Sul são unidades populacionais independentes e devem se tratar separadamente no planejamento e desenvolvimento de programas de manejo para a conservação da diversidade genética total da espécie.

碩士
臺灣大學
森林環境暨資源學研究所
96
Extra-Pair paternity (EPP) is defined as the offspring resulting from copulations outside the social pair bonds in the mating system. Field studies and comparative analyses have shown that EPP is due to female strategy. Females might obtain direct or indirect benefits through EPP. Female choice hypothesis predicts that females can pursue EPP and choose the preferred EPP males. By the contrary, EPP is constrained by males mate guarding behavior and females show no preference in the trade-off hypothesis. Taiwan yuhinas（Yuhina brunneiceps）is a joint-nesting, cooperatively breeding bird. The reproductive skew is low (0.19). Individuals in the breeding group share the load of nesting, the incubation and the provisioning. EPP has been recorded by observation in Taiwan yuhinas and there are 25.5% of EPP offspring have been recognized by paternity analysis. Data was selected from the breeding seasons during 2001-2006. Results showed that females prefer high-ranking males as EPP males and females might obtain indirect benefits from producing EPP offspring. Each member in the breeding group with its own demand and aim adapts different breeding strategies to get the highest reproductive benefits. Therefore, EPP is one kind of reproductive strategy in Taiwan yuhinas.

I investigated the role of extra-pair paternity on use of public information and the interaction between public information and personal information for patch fidelity decisions. It is unknown if songbirds use public information about the number of conspecific fledglings for patch fidelity decisions when extra-pair paternity is uncommon. I tested if probability of patch fidelity was associated with (1) number of fledglings in adjacent territories (public information), and (2) number of fledglings raised with a social mate (personal information). I used logistic regression to predict probability of patch fidelity of males and females based on the 2 uncorrelated predictor variables (Spearman’s rank correlation, S = 21895.28, n = 50, P = 0.723, r = –0.051). I monitored patch fidelity of 107 territories, counted the number of fledglings in each territory, and assessed parentage of 102 young from 36 nests for white-eyed vireos (Vireo griseus) from 2008–2010 in a 100 ha patch of woodland in central Texas, USA. I excluded the social male as the father of 3 of the 102 young and did not exclude any of the social females as the mother with parentage analysis using 6 microsatellite loci. The number of fledglings in adjacent territories was not a good predictor of probability of patch fidelity for males (beta 1 = 0.166, df = 35, P = 0.247, Nagelkerke’s R2 = 0.054) or females (beta 1 = 0.121, df = 17, P = 0.670, Nagelkerke’s R2 = 0.016). The number of fledglings raised with a social mate was also not a good predictor of probability of patch fidelity for males (beta 1 = –0.296, df = 43, P = 0.360, Nagelkerke’s R2 = 0.029), whereas it was a good predictor for females (beta 1 = 1.281, df = 21, P = 0.048, Nagelkerke’s R2 = 0.409). The dominant ecological concepts for explaining site fidelity in songbirds, win-stay lose-switch (based on personal reproductive success with a social mate) and public information, did not predict probability of patch fidelity well for male white-eyed vireos. The win-stay lose-switch model, but not public information was a good predictor of probability of patch fidelity for females. My results suggest that use of public information may depend on frequency of extra-pair paternity. Males may primarily use other information for patch fidelity decisions beyond reproductive success of conspecifics for patch fidelity decisions in some circumstances. My results support the need to ensure high levels of nesting success for females to return and maintain populations in areas managed for breeding songbirds for conservation efforts to be successful.

Molecular techniques have revealed that avian mating system is more diverse and complex than previously thought. Both males and females can use alternative reproductive tactics to increase their fitness. Here, we have determined the prevalence of conspecific brood parasitism (CBP, 22% of nests), quasi-parasitism (QP, 6.5% of nests) and extra-pair paternity (EPP, 51.2% of nests) in European subspecies of barn swallow (Hirundo rustica rustica). In contrast to EPP and CBP, QP is rare and has been described in only a few bird species. Our data indicate nonrandom QP patterns, suggesting that this tactic can be considered a third alternative reproductive strategy, alongside CBP and EPP. Sexual ornaments are usually assumed to evolve as condition-dependent signals of individual quality. In this context, a positive correlation between the expression of ornamental traits and survival, is expected. Evolutionary theory predicts that females seek extra-pair fertilizations from high-quality (more ornamented) males and EPP has consequently been proposed as a principal protagonist in the evolution of male secondary sexual traits, however empirical evidence in support of the assertion is relatively scarce. Despite decades of research into the role of EPP in sexual selection, the potential fitness advantages for...