Habitat selection and reproductive success of Ortolan Buntings Emberiza hortulana on farmland in central Sweden – the importance of habitat heterogeneity

Many granivorous birds have shown severe population declines in Europe during recent decades. The aim of the present study was to analyse habitat preferences and reproductive success of one such species, the Ortolan Bunting Emberiza hortulana, in different farmland habitats in south‐central Sweden. Four seemingly different land‐use types were preferred: permanent set‐asides, short rotation coppice, and grazed and unmanaged semi‐natural pastures. Territories and random sites differed considerably in the proportion of these preferred land‐use types; 39% of territories had > 70% preferred habitat (at the 100‐m scale) compared to 5% of random sites. In contrast, 22% of territories and 65% of random sites had no preferred habitats. All the preferred habitats had heterogeneous ground vegetation characterized by patches with bare ground, or at least sparse ground vegetation, intermixed with patches with taller vegetation. Ortolan Buntings also preferred a heterogeneous habitat structure with occurrence of field islets, shrubby edges, barns and electric wires, which could act as song posts or suitable nest‐sites, in 88% of territories. At a larger (1‐km square) scale, territories occupied by pairs aggregated strongly in areas with high proportions of preferred habitats. The number of territories with single males correlated positively with the number of pairs, which suggests that conspecific attraction may influence territory distribution. No measured habitat factors were related to reproductive success. However, due to habitat preferences and the higher proportion of paired males in one habitat type (set‐aside), the production of young (fledglings/ha) is expected to be higher in set‐asides, as well as in short‐rotation coppices and semi‐natural pastures. Thus, these habitats are important for the conservation of the Ortolan Bunting. Large areas with habitat structures such as field islets are especially important because the Ortolan Bunting breeds in aggregations in these areas.     

Sibling resemblance in natal dispersal distance and direction in the Ortolan Bunting Emberiza hortulana

Natal dispersal distance and direction determine the likelihood that siblings will settle close together and hence the risk of inbreeding. Several studies have shown a sibling resemblance in dispersal distance, but few studies have analysed sibling resemblance in dispersal direction or the distance between siblings after dispersal at the landscape level. I studied the entire Norwegian population of Ortolan Buntings Emberiza hortulana, which is patchily distributed in an area covering c. 500 km². Males and females did not differ in dispersal distance (overall median 3.7 km), but directions were different. Natal dispersal distances and directions were positively related within sibling pairs, but comparisons with control individuals suggested that any effects were due to spatial effects of configuration of habitat patches in the study area. Brother–sister pairs (n = 16) were at least as similar as brother–brother pairs (n = 18). Distance between siblings after natal dispersal increased with dispersal distance, but even so, five of 35 sibling pairs settled < 1 km apart, despite dispersal distances of 8.3–9.9 km for two of these pairs. Including movements later in life, eight sibling pairs were < 1 km apart at some time (four pairs of brothers and four brother–sister pairs), and in one case a brother mated with its sister. Another case of mating between close relatives (father and daughter) involved short female natal dispersal. These data indicate that female‐biased natal dispersal and long‐distance dispersal may reduce, but do not exclude, the possibility of inbreeding.     

Habitat selection by Ortolan Buntings Emberiza hortulana in post-fire succession in Catalonia: implications for the conservation of farmland populations

The Ortolan Bunting Emberiza hortulana is a long-distance migrant that has suffered major population declines across much of its European breeding range. While northern populations are bound largely to farmland, Mediterranean populations are largely confined to habitats subject to recurrent wildfires. Habitat selection of the Ortolan Bunting was assessed in a recently burnt area in Catalonia at landscape and habitat scales. A Zero-inflated Poisson procedure was used to model the abundance of birds in relation to landscape and habitat variables. The most parsimonious landscape model predicted the highest abundance on south-facing slopes, with a gradient above 10°. The most parsimonious habitat model showed a positive quadratic effect of bare ground and regenerating oak Quercus spp., with predicted optima for abundance around 20–30% and 20% cover, respectively. There was a clear relationship between predicted abundance of the Ortolan Bunting and post-fire regenerating oak shrubs. South-facing, moderately sloping areas were favoured and bare ground was a key feature of the species' habitat. A matrix combining patches of sparse oak shrubs and patches of bare ground appears to be the optimal breeding habitat in the Mediterranean. The maintenance or provision of similar habitat features, especially patches of bare ground, may prove crucial for the conservation of rapidly declining Ortolan Bunting populations on farmland across temperate Europe.     

Habitat preferences of the Ortolan Bunting (Emberiza hortulana) in its prime wintering grounds,the cereal-dominated Ethiopian Highlands

Agricultural intensification and land-use changes are major factors impacting farmland biodiversity. The Ortolan Bunting Emberiza hortulana is the long-distance trans-Saharan migratory passerine that has undergone the most dramatic decline among all European farmland birds. The factors responsible for this decline may originate from the breeding grounds, migration stopovers and/or overwintering quarters. Very little is known about conditions on the species' wintering grounds, but a recent study has highlighted the utmost importance of the traditionally managed agroecosystems in the Ethiopian Highlands as a key wintering area, apparently harbouring as much as 90% of the world's Ortolan Bunting population. Using radiotracking and line transect surveys, this study aimed to provide fine-grained information about species–habitat relationships in the Ortolan Bunting overwintering quarters. Our results showed the importance, at the landscape scale, of small-scale agriculture, notably of traditionally managed, cereal-dominated fields interspersed with semi-natural structures. At a foraging-site scale, on the other hand, patches of bare ground in combination with large areas of post-harvesting stubble represented key habitat features. Stubbles provide an essential food resource and bare ground promotes ground foraging by enhancing food accessibility. The maintenance of a traditional agricultural economy will be essential to maintain the habitat potential for the Ortolan Buntings overwintering in the Ethiopian Highlands and will be instrumental in preserving its world population from further decline.     

Lifetime fitness consequences of early‐life ecological hardship in a wild mammal population

Early‐life ecological conditions have major effects on survival and reproduction. Numerous studies in wild systems show fitness benefits of good quality early‐life ecological conditions (“silver‐spoon” effects). Recently, however, some studies have reported that poor‐quality early‐life ecological conditions are associated with later‐life fitness advantages and that the effect of early‐life conditions can be sex‐specific. Furthermore, few studies have investigated the effect of the variability of early‐life ecological conditions on later‐life fitness. Here, we test how the mean and variability of early‐life ecological conditions affect the longevity and reproduction of males and females using 14 years of data on wild banded mongooses (Mungos mungo). Males that experienced highly variable ecological conditions during development lived longer and had greater lifetime fitness, while those that experienced poor early‐life conditions lived longer but at a cost of reduced fertility. In females, there were no such effects. Our study suggests that exposure to more variable environments in early life can result in lifetime fitness benefits, whereas differences in the mean early‐life conditions experienced mediate a life‐history trade‐off between survival and reproduction. It also demonstrates how early‐life ecological conditions can produce different selection pressures on males and females.     

Lifetime reproductive success,selection on lifespan,and multiple sexual ornaments in male European barn swallows

Natural and sexual selection arise when individual fitness varies according to focal traits. Extra‐pair paternities (EPPs) can affect the intensity of selection by influencing variance in fitness among individuals. Studies of selection require that individual fitness is estimated using proxies of lifetime reproductive success (LRS). However, estimating LRS is difficult in large, open populations where EPPs cause reallocation of biological paternity. Here, we used extensive field sampling to estimate LRS in a population of barn swallows (Hirundo rustica) to estimate selection on lifespan and ornamental traits of males. We found selection on lifespan mediated both by within‐ and extra‐pair fertilization success and selection on tail length mediated by within‐ but not extra‐pair fertilization success. In addition, we found selection on tail white spots via extra‐pair fertilization success after controlling for selection on other traits. These results were not confounded by factors that hamper studies of LRS, including nonexhaustive sampling of offspring and biased sampling of males. Hence, natural and sexual selection mediated by LRS operates on lifespan, tail length, and size of the tail white spots in barn swallows.     

Lifetime coastal and oceanic foraging patterns of male Guadalupe fur seals and California sea lions

Carcasses of adult and subadult male Guadalupe fur seals (GFSs) are beached on Isla Magdalena, representing about 73% of all GFSs beachings since 2003. Analysis of carbon and nitrogen stable isotopes in dentin collagen from male GFSs was used in an attempt to distinguish animals from two possible locations of origin: the oceanic Guadalupe Island (GI) and the coastal San Benito Islands (SBI). Samples of conterminous and contemporary California sea lions (CSLs) were included as representative of coastal predators. Two distinct groups of GFSs were evident based on δ¹³C values: coastal GFSs with higher δ¹³C values than CSLs, and oceanic GFSs with lower δ¹³C values than CSLs. Significant temporal trends were observed in the δ¹⁵N values; values for oceanic GFSs and CSLs increased annually (0.082‰ and 0.089‰, respectively), while those for coastal GFS decreased annually by 0.183‰. The isotopic segregation of oceanic and coastal GFS males, presumably from GI and SBI respectively, has been present at least since the mid-1990s. This segregation may be due to dissimilar ingestion of prey from oceanic vs. coastal origin prior to their seasonal co-occurrence in the Gulf of Ulloa.     

Determinants of age at first reproduction and lifetime breeding success revealed by full paternity assignment in a male ungulate

Age at first reproduction is an important determinant of individual variation in reproductive success in ungulates, but few studies have examined its relationship with later fitness‐related traits in males. We used a long‐term individual based study of a harvested moose population to quantify the individual reproductive performance and survival of males, as well as to examine the determinants of age at first reproduction and consequences of age at first reproduction on lifetime breeding success. The probability that a male successfully reproduced at the age of two was negatively related to the mean age of adult males in the population, but the relationship weakened with increasing population size. Large antlers and large body mass relative to other males in the population increased the number of calves sired at their first successful mating season. In addition, those that successfully reproduced as two year‐olds were more likely to sire calves the next year, making them more productive at a given age compared to those that first reproduced at the age of three or older. We emphasize the importance for males to start reproducing as soon as possible in a harvested population to gain lifetime fitness benefits, as surviving the hunt is a major determinant of reproductive success in this population. We found no costs of early reproduction in males, hence leading to high individual heterogeneity in male reproductive performance. The apparent lack of reproductive costs could partly be explained by the age distribution in the population, individual variation in early‐life body mass and antler size, and differences in probabilities of being hunted of successful and unsuccessful males.     

Influence of fluctuation in the operational sex ratio to mating of troop and non-troop male Japanese macaques for four years on Kinkazan Island, Japan

Taking advantage of a marked yearly fluctuation in the number of estrous females, I studied the differences in mating success between troop males and non-troop males in an unprovisioned group of Japanese macaques. Fluctuation in the defendability of estrous females by troop males, as predicted by the operational sex ratio (the number of estrous females per troop male), strongly affected the mating with ejaculation (successful mating) per observation day of both troop and non-troop males. When operational sex ratio was low, troop males monopolized successful mating inside the troop. No successful mating of non-troop males was observed inside the troop. In contrast, both troop and non-troop males were able to mate often inside the troop when operational sex ratio was high. These findings suggest that troop males obtained the benefit of secured successful mating in the troop because troop males could mate successfully even in mating seasons with a low operational sex ratio, and the chance of successful mating for non-troop males will increase as the ability of troop males to monopolize estrous females decreases.     

Culled males,infant mortality and reproductive success in a pre-industrial Finnish population

Theoretical and empirical literature asserts that the sex ratio (i.e. M/F) at birth gauges the strength of selection in utero and cohort quality of males that survive to birth. We report the first individual-level test in humans, using detailed life-history data, of the ‘culled cohort’ hypothesis that males born to low annual sex ratio cohorts show lower than expected infant mortality and greater than expected lifetime reproductive success. We applied time-series and structural equation methods to a unique multigenerational dataset of a natural fertility population in nineteenth century Finland. We find that, consistent with culled cohorts, a 1 s.d. decline in the annual cohort sex ratio precedes an 8% decrease in the risk of male infant mortality. Males born to lower cohort sex ratios also successfully raised 4% more offspring to reproductive age than did males born to higher cohort sex ratios. The offspring result, however, falls just outside conventional levels of statistical significance. In historical Finland, the cohort sex ratio gauges selection against males in utero and predicts male infant mortality. The reproductive success findings, however, provide weak support for an evolutionarily adaptive explanation of male culling in utero.     

The influence of female age on male mating preference and reproductive success in cabbage beetle,Colaphellus bowringi

The influence of female age on male mating preference and reproductive success has been studied using a promiscuous cabbage beetle, Colaphellus bowringi Baly （Coleoptera： Chrysomelidae）. In a simultaneous choice test, middle-aged females had significantly greater mating success than young and old females. In single pair trials, when paired with middle-aged virgin males, middle-aged females mated faster, copulated longer, and had greater fecundity and fertility than young or old females, while the longevity of males was not significantly affected by female age. This study on C. bowringi suggests that middle-aged females are more receptive to mating, which can result in the highest male reproductive success.     

豆野螟成虫日龄对交尾的影响及雄蛾对性信息素的触角电位反应

豆野螟Marucavitrata(Fabricius)是一种严重的泛热带豆类蔬菜害虫。在不同温度,光周期14L∶10D,相对湿度75%～80%条件下,研究豆野螟成虫日龄对交尾的影响及雄蛾对性信息素的EAG反应。不同日龄雌蛾与3日龄雄蛾的交尾,交尾率随蛾龄增加呈下降趋势;不同日龄雄蛾与3日龄雌蛾交尾,交尾率随蛾龄增加呈升高趋势;相同日龄成蛾,交尾率先升后降。不同日龄雌蛾交尾百分率的下降在较高温环境比在较低温环境更加显著;但环境温度对不同日龄雄蛾的交尾却没有太大的影响;相同日龄成蛾3日龄以后的交尾在较高温环境下下降的更快。不同日龄雌蛾的交尾研究中,随着雌蛾日龄的增加,交尾持续时间明显延长;不同日龄雄蛾的交尾研究中,较低日龄和较高日龄雄蛾交尾时的持续时间较长。在较低温环境下所有试验各日龄的持续时间均有所延长。不同日龄雌蛾(雄蛾)与3日龄雄蛾(雌蛾)的交尾和相对应的雄蛾对雌蛾性信息素的EAG反应基本一致,说明雄蛾的反应在交尾行为的完成中具有重要的作用。     

Density-dependent self-fertilization and male versus hermaphrodite siring success in an androdioecious plant

Models of mating-system evolution emphasize the importance of frequency-dependent interactions among mating partners. It is also known that outcross siring success and the selfing rate in self-compatible hermaphrodites can be density dependent. Here, we use array experiments to show that the mating system (i.e., the outcrossing rate) and the siring success of morphs with divergent sex allocation strategies are both density dependent and frequency dependent in androdioecious populations of the wind-pollinated, annual plant Mercurialis annua. In particular, the outcrossing rate is a decreasing function of the mean interplant distance, regulated by a negative exponential pollen fall-off curve. Our results indicate that pollen dispersed from a male inflorescence are over 60% more likely to sire outcrossed progeny than equivalent pollen dispersed from hermaphrodites, likely due to the fact that males, but not hermaphrodites, disperse their pollen from erect inflorescence stalks. Because of this difference, and because males of M. annua produce much more pollen than hermaphrodites, the presence of males in the experimental arrays reduced both the selfing rate and the outcross siring success of hermaphrodites. We use our results to infer a density threshold below which males are unable to persist with hermaphrodites but above which they can invade hermaphroditic populations. We discuss our findings in the context of a metapopulation model, in which males can only persist in well-established populations but are excluded from small, sparse populations, for example, in the early stages of colonization.     

Sex-specific sibling interactions and offspring fitness in vertebrates: patterns and implications for maternal sex ratios

Vertebrate sex ratios are notorious for their lack of fit to theoretical models, both with respect to the direction and the magnitude of the sex ratio adjustment. The reasons for this are likely to be linked to simplifying assumptions regarding vertebrate life histories. More specifically, if the sex ratio adjustment itself influences offspring fitness, due to sex-specific interactions among offspring, this could affect optimal sex ratios. A review of the literature suggests that sex-specific sibling interactions in vertebrates result from three major causes: (i) sex asymmetries in competitive ability, for example due to sexual dimorphism, (ii) sex-specific cooperation or helping, and (iii) sex asymmetries in non-competitive interactions, for example steroid leakage between fetuses. Incorporating sex-specific sibling interactions into a sex ratio model shows that they will affect maternal sex ratio strategies and, under some conditions, can repress other selection pressures for sex ratio adjustment. Furthermore, sex-specific interactions could also explain patterns of within-brood sex ratio (e.g. in relation to laying order). Failure to take sex-specific sibling interactions into account could partly explain the lack of sex ratio adjustment in accordance with theoretical expectations in vertebrates, and differences among taxa in sex-specific sibling interactions generate predictions for comparative and experimental studies.     

Ritual fights and male reproductive success in a human population

Ritual fights are widespread across human populations. However, the evolutionary advantage associated with this behaviour is unclear because these fights rarely provide direct benefits such as territory, resources or mates. Here, the reproductive success of men competing in a traditional ritual fight, Sereer wrestling, was investigated for the first time. Involvement in wrestling had a significant positive effect on men’s number of offspring and a marginally significant effect on polygyny, controlling for age, body condition and socio‐economic status. These positive effects suggest that being involved in wrestling competition provides prestige, facilitating access to mates and thereby increasing fecundity. However, when women were interviewed on their preference concerning qualities of potential mates, the quality ‘being involved in wrestling competition’ was poorly ranked. This discrepancy may arise either from deceptive reports or from discordance between parents and daughters in the choice of a husband.     

Lifetime reproductive success and longevity of queens in an annual social insect

Although central to understanding life‐history evolution, the relationship between lifetime reproductive success and longevity remains uncertain in many organisms. In social insects, no studies have reported estimates of queens’ lifetime reproductive success and longevity within populations, despite the importance of understanding how sociality and associated within‐group conflict affect life‐history traits. To address this issue, we studied two samples of colonies of the annual bumblebee, Bombus terrestris audax, reared from wild‐caught queens from a single population. In both samples, queens’ lifetime reproductive success, measured as either queens’ inclusive fitness or as total biomass of queen‐produced sexuals (new queens and males), was significantly positively associated with queen longevity, measured from the day the first worker was produced. We suggest that a positive relationship between reproductive success and longevity was inherited from nonsocial ancestors showing parental care and maintained, at least in part, because the presence of workers buffers queens against extrinsic mortality.     

One-male units and clans in a colony of hamadryas baboons (Papio hamadryas hamadryas): effect of male number and clan cohesion on feeding success

In the multilevel societies of hamadryas baboons, adult males can be attached to single one-male units (OMUs) or to clans containing several such OMUs. This paper examines the effect of male number and rivalry between males within a clan on their ability to compete for access to a clumped food resource. The data come from a study of a multilevel colony of hamadryas baboons (Papio hamadryas hamadryas) housed at the Madrid Zoo. The colony consisted of 12 harem-holding males and 40 sexually mature females, and was organized into five single OMUs and two clans (containing three and four OMUs, respectively). The top-ranking male of one of the clans was removed and later reintroduced, so the study involved an analysis of the composition of clans and OMUs and of the males' use of the feeding area across three study periods: preseparation, separation, and reintroduction. The findings reported indicate that both males and females derived clear advantages in the context of contest competition for access to clumped food if they were members of clans, because the males and females from large clans had a feeding advantage over those from smaller clans and single OMUs. Furthermore, rivalry among males within the clan reduced their ability to compete for food against males outside their clan. This paper provides empirical evidence for one of the potential advantages that hamadryas males may enjoy if they are attached to clans, and also provides empirical support for the general hypothesis that a large number of males in a group may provide fitness-related benefits to the group members, provided they are able to cooperate with each other.