Philopatry and natal dispersal of Montagu's harriers (Circus pygargus) breeding in Spain: a review of existing data

Natal dispersal is an important component of bird ecology, plays a key role in many ecological and evolutionary processes, and has important conservation implications. Nevertheless, detailed knowledge on natal dispersal is still lacking in many bird species, especially raptors. We review and compile existing information from five tagging programmes of juvenile Montagu's harriers (Circus pygargus) in different Spanish regions, with PVC rings or wing tags, to provide an assessment of philopatry and natal dispersal of the species in Spain. Only 7% of all tagged harriers were observed as breeders in subsequent years. The percentage of philopatric (i.e. breeding within 10?km of the natal site) males and females was lower that 5%. Overall, there were no sexual differences in percentage of philopatric birds or dispersal distances, but we found study area differences. The low philopatry observed suggests a high capacity for natal dispersal in this species, for both sexes, and therefore high genetic mixing between populations. Differences in philopatry between study areas may be influenced by the different observation effort or detectability, or else reflect different philopatric strategies among populations. Finally, we found no significant differences in philopatry rate or dispersal distances related to tagging method, suggesting that tagging technique has a smaller effect than monitoring effort or observation ease on observation probability. Developing tagging programmes at a small scale and without procuring very large-scale and intensive subsequent monitoring is not worthwhile for evaluating philopatry and natal dispersal in this species.     

Social dominance in free-living Willow Tits Parus montanus: determinants and some implications of hierarchy

Several features of social dominance among Willow Tit Parus montanus winter flocks were examined during a four-winter study. Birds of both sexes were evenly distributed over the 33 flocks studied. In nearly half of the flocks there was an adult pair accompanied by yearlings, but one-third of the flocks consisted of more than two adults with yearlings. Males were found to be dominant over and larger than females. Within a sex, yearlings were usually subordinate to adults. The effect of size on dominance, after controlled for sex and age, remained obscure in our field data. The hierarchical status of an individual was found to rise or at least stay the same in different years, which supports the "hopeful dominants" hypothesis, i.e. birds stay in flocks hoping to achieve a higher status in the future. The ranks of mates correlated highly significantly, implying that high-ranking birds were paired with other high-ranking birds and low-ranking birds with other low-ranking birds. Birds of different age and sex did not show any differences in the proportion of initiated aggressive encounters directed at other individuals. However, males were more aggressive to other males than to females and also tended to behave less aggressively towards their own mates than towards other individuals in the flock. This could be a male strategy to strengthen the pair-bond and to enhance mate protection described earlier in     

Genetic evidence for sex-biased dispersal in resident bottlenose dolphins (Tursiops aduncus)

In most mammals males usually disperse before breeding, while females remain in their natal group or area. However, in odontocete cetaceans behavioural and/or genetic evidence from populations of four species indicate that both males and females remain in their natal group or site. For coastal resident bottlenose dolphins field data suggest that both sexes are philopatric to their natal site. Assignment tests and analyses of relatedness based on microsatellite markers were used to investigate this hypothesis in resident bottlenose dolphins, Tursiops aduncus, from two small coastal populations of southeastern Australia. Mean corrected assignment and mean relatedness were higher for resident females than for resident males. Only 8% of resident females had a lower probability than average of being born locally compared to 33% of resident males. Our genetic data contradict the hypothesis of bisexual philopatry to natal site and suggest that these bottlenose dolphins are not unusual amongst mammals, with females being the more philopatric and males the more dispersing sex.     

The spatial distribution of avian relatives: do obligate army‐ant‐following birds roost and feed near family members?

The ocellated antbird ( Phaenostictus mcleannani ) feeds in groups and therefore is an informative species in which to study the biological factors that modulate avian group living. These birds congregate at swarms of army ants to capture fleeing prey, and previous observations suggest that males may be philopatric, feed with close relatives, and defend communal feeding ranges. We assessed whether kin selection could be an important factor maintaining group formation in a population of ocellated antbirds inhabiting continuous forest at La Selva Biological Station, Costa Rica, using radiotelemetry and 15 novel microsatellite markers. We predicted that the roosting areas of closely related adult males should overlap and that adult males feeding simultaneously at the same swarm should be highly related. We banded and genotyped 65 individuals (≥ 88% of the population) and radiotagged 30 of them. The results generally did not conform to our predictions. Little overlap occurred among the roosting areas of same-sex individuals, and nearest roosting neighbours (either same or opposite sex) were generally unrelated. A small proportion of male dyads suggested short-distance dispersal, but in general the distribution of genotypes within the study area approached randomness. We found little evidence of natal philopatry in either sex. Less than half of the feeding groups sampled included highly related males; most consisted of unrelated individuals. Hence, we found limited potential for kin selection to favour group living and suggest that other factors, particularly direct benefits (e.g. food intake), are probably more important than indirect effects (nepotism).     

Philopatry, Site Fidelity and Local Kin Associations within Grey Seal Breeding Colonies

Abstract Colonially breeding animals such as grey seals typically have a discrete number of breeding sites. The extent to which offspring are philopatric (return to breed at their natal site) and the prevalence of philopatry by sex of pup has fundamental consequences for the social and genetic structure within and between each colony.
Grey seals born and marked by tagging and cohort brands at colonies on North Rona, Outer Hebrides since 1960 and at the Isle of May, Firth of Forth since 1981 have shown philopatry. Overall re-sight rates of 1620 pups tagged at North Rona and 1667 pups at the Isle of May ranged from 0 to 17%. Although most evidence of philopatry relates to females, males at each colony displayed philopatry. In addition, females born at N. Rona pupped closer to their natal sites than would be expected by chance (p = 0.005), providing evidence of fine scale natal site fidelity. Females born at the Isle of May did not show the same degree of fine scale natal site fidelity, but there was considerable individual variation. The spatial scale of philopatry shown by seals differed according to the local area within each island. Fidelity of known adult females to pupping sites at both colonies was high. In addition, we found an instance of mother and daughter pupping together away from the daughter's natal site.
These data: (i) provide evidence of philopatry in both sexes at each site; (ii) describe occurrences of fine scale philopatry at both sites; (iii) show that mothers and offspring occupied the same areas on the colony, suggesting the possibility of highly related groups within colonies, which would provide conditions for kin-specific altruistic behaviours; (iv) suggest the possibility of individual or kin recognition, even though contact between mothers and offspring has been thought to cease after the 18 d lactation period.     

Elevated mortality among birds in Chernobyl as judged from skewed age and sex ratios

Background

Radiation has negative effects on survival of animals including humans, although the generality of this claim is poorly documented under low-dose field conditions. Because females may suffer disproportionately from the effects of radiation on survival due to differences in sex roles during reproduction, radiation-induced mortality may result in male-skewed adult sex ratios.

Methodology/Principal Finding

We estimated the effects of low-dose radiation on adult survival rates in birds by determining age ratios of adults captured in mist nets during the breeding season in relation to background radiation levels around Chernobyl and in nearby uncontaminated control areas. Age ratios were skewed towards yearlings, especially in the most contaminated areas, implying that adult survival rates were reduced in contaminated areas, and that populations in such areas could only be maintained through immigration from nearby uncontaminated areas. Differential mortality in females resulted in a strongly male-skewed sex ratio in the most contaminated areas. In addition, males sang disproportionately commonly in the most contaminated areas where the sex ratio was male skewed presumably because males had difficulty finding and acquiring mates when females were rare. The results were not caused by permanent emigration by females from the most contaminated areas because none of the recaptured birds had changed breeding site, and the proportion of individuals with morphological abnormalities did not differ significantly between the sexes for areas with normal and higher levels of contamination.

Conclusions/Significance

These findings are consistent with the hypothesis that the adult survival rate of female birds is particularly susceptible to the effects of low-dose radiation, resulting in male skewed sex ratios at high levels of radiation. Such skewed age ratios towards yearlings in contaminated areas are consistent with the hypothesis that an area exceeding 30,000 km² in Chernobyl’s surroundings constitutes an ecological trap that causes dramatic excess mortality.     

Sex-biased dispersal and natal philopatry in the diamondback terrapin, Malaclemys terrapin

Nesting ecology and population studies indicate that diamondback terrapins (Malaclemys terrapin) exhibit nest site fidelity and high habitat fidelity. However, genetic studies indicate high levels of gene flow. Because dispersal affects the genetics and population dynamics of a species, we used six highly polymorphic microsatellite markers to investigate sex-biased dispersal and natal philopatry of M. terrapin in Barnegat Bay, NJ. We compared results of spatial autocorrelation analysis, assignment methods and Wright's F(ST) estimators to a mark-recapture analysis. Mark-recapture analysis over a 4-year period indicated that most individuals have relatively small home ranges (<2 km), with mature females displaying greater home ranges than males. Goodness of fit analysis of our mark-recapture study indicated that some juvenile males were likely transient individuals moving through our study location. Mean assignment indices and first-generation migrant tests indicated that mature males were more prone to disperse than mature females, but first-generation migrant tests indicated that per capita there are more female than male dispersers. Thus, the relative importance of males and females on gene flow in terrapin populations may change in relation to population sex ratios. Spatial autocorrelation analysis indicated that mature females exhibited natal philopatry to nesting beaches, but first-generation migrant tests indicated that a small number of females failed to nest on natal beaches. Finally, we discuss the important conservation implications of male-biased dispersal and natal philopatry in the diamondback terrapin.     

Female‐biased dispersal in a bat with a female‐defence mating strategy

The ultimate causes for predominant male‐biased dispersal (MBD) in mammals and female‐biased dispersal (FBD) in birds are still subject to much debate. Studying exceptions to general patterns of dispersal, for example, FBD in mammals, provides a valuable opportunity to test the validity of proposed evolutionary pressures. We used long‐term behavioural and genetic data on individually banded Proboscis bats (Rhynchonycteris naso) to show that this species is one of the rare mammalian exceptions with FBD. Our results suggest that all females disperse from their natal colonies prior to first reproduction and that a substantial proportion of males are philopatric and reproduce in their natal colonies, although male immigration has also been detected. The age of females at first conception falls below the tenure of males, suggesting that females disperse to avoid father–daughter inbreeding. Male philopatry in this species is intriguing because Proboscis bats do not share the usual mammalian correlates (i.e. resource‐defence polygyny and/or kin cooperation) of male philopatry. They have a mating strategy based on female defence, where local mate competition between male kin is supposedly severe and should prevent the evolution of male philopatry. However, in contrast to immigrant males, philopatric males may profit from acquaintance with the natal foraging grounds and may be able to attain dominance easier and/or earlier in life. Our results on Proboscis bats lent additional support to the importance of inbreeding avoidance in shaping sex‐biased dispersal patterns and suggest that resource defence by males or kin cooperation cannot fully explain the evolution of male philopatry in mammals.     

Philopatry and within-colony movements in Columbian ground squirrels

Philopatry and dispersal result in selection of habitat locations that may differ in resources and social environment and thus should influence fitness components like survival and reproduction. We examined short-distance movements of young and adult females from natal or previous nesting sites within a colony of Columbian ground squirrels (Urocitellus columbianus) in the Rocky Mountains of Alberta, Canada, over a 17-year period. Females of all ages were strongly philopatric, yet a few (10-15%) exhibited movements that took them to new home ranges. We tested three hypotheses to explain the pattern of female natal and breeding movements: (1) that movements of philopatric females promote proximity to close kin; (2) that range shifts favour close kin via bequeathal of territory and (3) that dispersers move to lower density areas where competition for resources is lower. Tests of these three hypotheses revealed that: (1) philopatry and movements of young and older philopatric females led to proximity to mothers and local presence of close kin; (2) breeding dispersal did not result in bequeathal of home range to daughters, but movements of philopatric females suggested that they shared space with close kin and (3) adult females moved to new ranges with lower local densities, though dispersing females also left ranges where local density was significantly lower than for philopatric females. Natal and breeding movements among years produced two opportunities for territorial females: close spatial proximity to close kin via short philopatric movements, and habitats with fewer competitors via longer dispersal movements.     

Dispersal costs set the scene for helping in an atypical avian cooperative breeder

The ecological constraints hypothesis is suggested to explain the evolution of cooperative breeding in birds. This hypothesis predicts that the scene for cooperative breeding is set when ecological factors constrain offspring from dispersal. This prediction was tested in the atypical cooperative breeding system of the long-tailed tit, Aegithalos caudatus, by comparing the degree of philopatry and cooperation in an isolated and a contiguous site whilst experimentally controlling for confounding aspects of reproduction. No difference was found between the two sites in the survival of offspring but a greater proportion were found to remain philopatric in the isolated site. This difference was caused by greater philopatry of normally dispersive females suggesting, as predicted, that dispersal costs were greater from this site. Furthermore, a greater proportion of males and females cooperated following breeding failure in the isolated site than in the contiguous site. Thus, as has been suggested for typical avian cooperative breeders, dispersal costs, relative to philopatric benefits, appear to set the scene for cooperative breeding in long-tailed tits.     

Mating systems,philopatry and dispersal in birds and mammals

Many species of birds and mammals are faithful to their natal and breeding site or group. In most of them one sex is more philopatric than the other. In birds it is usually females which disperse more than males; in mammals it is usually males which disperse more than females. Reproductive enhancement through increased access to mates or resources and the avoidance of inbreeding are important in promoting sex differences in dispersal. It is argued that the direction of the sex bias is a consequence of the type of mating system. Philopatry will favour the evolution of cooperative traits between members of the sedentary sex. Disruptive acts will be a feature of dispersers.     

Age differences in blue tit Parus caeruleus plumage colour: within‐individual changes or colour‐biased survival?

In many species of passerine birds yearlings display a less elaborate version of the adult secondary sexual traits, but the causes of such differences in ornamentation are not always well understood. We studied age-related changes in blue tit Parus caeruleus UV/blue structural crown coloration, a sexually selected trait. In our Austrian study population, older blue tits, irrespective of sex, displayed on average a more ultraviolet (lower hue, higher UV chroma), more chromatic and brighter crown coloration than yearlings. This age dichromatism was caused by within-individual changes in the expression of crown coloration between years since males and females became more UV, more chromatic and brighter as they aged. Colour biased survival did not contribute to the observed pattern of age dichromatism since crown coloration was largely unrelated to overwinter survival. Between-year repeatability of crown colour was significant for most colour variables but low in general, and lower for females than for males. In the blue tit, yearling males might benefit from being less ornamented by avoiding adult aggression but at the expense of sexual attractiveness. Adaptive explanations of blue tit age dichromatism should however take into account that age effects were of similar magnitude in males and females. This suggests that both male and female yearlings could benefit from being less ornamented and hence that sexual selection might be acting on both sexes simultaneously in this species.     

Sex-biased dispersal: a result of a sex difference in breeding site availability

Sex-biased dispersal is often explained by assuming that the resource-defending sex pays greater costs of moving from a familiar area. We hypothesize that sex-biased dispersal may also be caused by a sex bias in breeding site availability. In avian resource-defense mating systems, site availability is often more constrained for females: males can choose from all vacant sites, whereas females are restricted to sites defended by males. Using data on breeding dispersal of a migratory passerine, we show that average number of available breeding options and availability of the previous year's territory was greater for males than females. The female bias in site unavailability may explain the female bias in probability of breeding dispersal because there was no sex bias in dispersal among birds with their previous year's territory available. We suggest that sex biases in the availability of breeding options may be an important factor contributing to observed variation in sex-biased dispersal patterns.     

Identification and breeding of yearling Piping Plovers

ABSTRACT Age of first breeding is an important life history trait. Many Piping Plovers (Charadrius melodus) do not breed as yearlings, but little information is available concerning the age of first breeding. From 2000 to 2006, we marked 991 chicks in three areas in Saskatchewan, Canada, and subsequently determined when 102 (49 females and 53 males) first bred. Females bred significantly earlier, on average, than males. More females (68%) bred as yearlings than did males (41%; P= 0.04), with most others first nesting when 2‐yr old (29% of females and 50% of males). As expected from differences between the sexes in age of first breeding, younger females were more likely to pair with older males than were younger males with older females. Chicks that hatched early in the breeding season did not breed at an earlier age than those that hatched late in the year. Unlike older birds, juvenile Piping Plovers do not replace flight feathers during their first winter. As a result, 18 of 27 yearlings (67%) had worn outer primaries, whereas only one of 123 (1%) older birds had worn primaries. In addition, whereas 20 of 24 yearlings (83%) retained a few buff‐tipped median coverts, none of 119 known older birds had such coverts. As a result, we were able to identify all yearlings by their worn primaries, buff‐tipped median wing coverts, or both. Wing lengths of yearling Piping Plovers were 3% shorter than those of older birds, presumably due to wear. Because there is no evidence of differences in adult survival rates between the sexes and breeding habitat is available, we speculate that fewer yearling males than females breed because primary wear may reduce the ability of yearling males to perform aerial breeding displays.     

Should females of migratory dabbling ducks switch mates between wintering and breeding sites?

Pairing of northern hemisphere dabbling ducks normally occurs in wintering sites. Mate choice occurs when some individuals congregate in courtship parties, in which females mainly choose mates according to male behavioural dominance. The sex ratio of some species is more male-biased at northern than at southern wintering sites, and the age ratio is adult-biased in northern areas. A possible mechanism responsible for this spatial segregation of sex and age classes is behavioural dominance, with males usually dominating females and adults dominating yearlings. Due to this latitudinal segregation of sexes during winter, presumably more males would remain unpaired in northern sites utilized by the more dominant birds than in southern sites utilized by the less-dominant birds, and this is paradoxical. I argue that some females having wintered and paired in southern sites may switch mates at stopover sites during northward migration. This hypothesis provides an explanation as to why, in spite of females being paired when initiating northward migration, some species show an increase in courtship activity during spring at stopover sites, as in common teals (Anas crecca) in southern Spain. In contrast, populations that do not migrate, e.g. mallards (A. platyrhynchos) in southern Spain, do not exhibit an increase in courtship activity during spring.     

Variation in Survival and Harvest Rates in Florida Mottled Duck

The Florida mottled duck (Anas fulvigula fulvigula) inhabits a relatively small range of approximately 90,000 km² within peninsular Florida, USA, and is threatened by habitat loss and genetic introgression with feral mallards (Anas platyrhynchos). Moreover, the Florida mottled duck population status has not been assessed for more than a decade. We used band-recovery and recapture data from 2000–2013 to examine geographic and demographic factors that influence the survival of Florida mottled ducks and to determine whether survival and harvest probabilities have changed over time. Mean survival probabilities were higher for birds banded in the southern portion of their Florida range than for those banded in the northern portion and higher for adult males than for adult females in both areas. Harvest probabilities increased in the northern extent of its range in Florida for adults and juveniles and remained relatively constant in the southern portion of its range during the study period. Mean harvest probabilities for adult males in both areas were higher than for adult females. Mean harvest probability for juvenile females was higher than that for juvenile males in the north but was similar between the sexes in the south. Our results suggest that mortality rates are generally greater in the northern portion of the Florida mottled duck range because of regional differences in habitat distribution and permanence and in how mottled ducks and humans use wetlands in these areas. We suggest increasing conservation efforts in the north portion of the Florida mottled duck range and improving inferences from leg banding by incorporating live recapture data. © 2020 The Wildlife Society.     

Male‐skewed adult sex ratio,survival, mating opportunity and annual productivity in the Snowy Plover Charadrius alexandrinus

Sex differences in adult mortality may be responsible for male‐skewed adult sex ratios and male‐skewed parental care in some birds. Because a surplus of breeding males has been reported in serially polyandrous populations of Snowy Plover Charadrius alexandrinus, we examined sex ratio, early‐season nesting opportunities, adult survival and annual reproductive success of a Snowy Plover population at Monterey Bay, California. We tested the hypotheses that male adult survival was greater than female survival and that a sex difference in adult survival led to a skewed adult sex ratio, different mating opportunities and different annual productivity between the sexes. Virtually all females left chicks from their first broods to the care of the male and re‐nested with a new mate. As a result, females had time to parent three successful nesting attempts during the lengthy breeding season, whereas males had time for only two successful attempts. Among years, the median population of nesting Plovers was 96 males and 84 females (median difference = 9), resulting in one extra male per eight pairs. The number of potential breeders without mates during the early nesting period each year was higher in males than in females. Adult male survival (0.734 ± 0.028 se) was higher than female survival (0.693 ± 0.030 se) in top‐ranked models. Annually, females parented more successful clutches and fledged more chicks than their first mates of the season. Our results suggest that in C. alexandrinus a sex difference in adult survival results in a male‐skewed sex ratio, which creates more nesting opportunities and greater annual productivity for females than for males.     

Sex differences in the stopover ecology of Curlew Sandpipers Calidris ferruginea at a refuelling area during autumn migration

We investigated the stopover patterns of male and female Curlew Sandpipers at a stopover area in northeast Spain. Curlew Sandpipers were trapped and colour-ringed during autumn migration in 1992 and 1993. Stopover length was similar to those reported previously for this and other waders that migrate using a small number of widely separated staging areas, but were greater than stopovers reported for other waders that migrate using a large number of staging areas separated by short distances. The differences in stopover length between the birds using these two strategies could be related to the fuel reserves that have to be accumulated to reach the next staging area. Males stayed longer in the area than females. Seasonal changes in prey availability or sex differences in moulting and migratory patterns do not account for these differences in stopover ecology. Following a time-selected model of optimal migration, sex differences in stopover ecology could be related to a dominance of the larger females over the males or to a higher foraging efficiency or a shorter search and settling time in females. Whether these differences are restricted to the studied area or are widespread in other staging areas used by the species could be important for assessing the possible differences in the migration speed of Curlew Sandpipers. The finding that males leave the breeding grounds 21–35 days before females but arrive at the study area with only a 10-day difference supports the hypothesis that females migrate faster than males at least in the first half of their migration.     

Ecological Correlates of Roost Fidelity in the Tent-Making Bat Artibeus watsoni

Roost switching is a common occurrence in bats, yet the causes and consequences of such behavior are poorly understood. In this study we explore the ecological correlates of roost fidelity in the tent‐making bat Artibeus watsoni, particularly focusing on the effect of sex, reproductive status, and roost availability using a three‐factor general linear model (GLM). We estimated roost fidelity of radio‐tracked individuals and found that the GLM was significant (R² = 0.72, F_10,34 = 8.91, p < 0.001). Significant interaction terms were observed for relative roost availability and sex (F_4,34 = 16.96, p < 0.001), and relative roost availability and reproductive status (F_6,34 = 7.62, p < 0.001), indicating that variation in roost fidelity among males and females, and among individuals under different breeding conditions, depended on relative roost availability at the site where they were radio‐tracked. Individuals in areas of high roost availability exhibited lower roost fidelity than those sampled in areas of lower roost availability. Females exhibited less roost fidelity than males for all roost availability categories, but the difference between males and females was only significant at high roost availability. The general pattern of decreased roost fidelity as roost availability increased was also prevalent among individuals in different breeding conditions. Additionally, satellite males exhibited higher roost fidelity than resident males in areas of low roost availability, and lactating females had higher roost fidelity than non‐breeding females in areas of medium roost availability. Our study thus demonstrates that sex, reproductive status, and roost availability all affect roost fidelity in the tent‐making bat A. watsoni, and also suggests that roost availability is the most important factor influencing roost fidelity in this bat, providing the first quantitative evidence that roost fidelity is correlated with roost abundance in a single species.