Dispersal, interpatch movements, and survival in a shrubland breeding bird community

ABSTRACT Dispersal events can affect the distribution, abundance, population structure, and gene flow of animal populations, but little is known about long‐distance movements due to the difficulty of tracking individuals across space. We documented the natal and breeding dispersal of shrubland birds among 13 study sites in a 1000 km² area in southeastern Ohio. In addition, we radio‐marked and tracked 37 adult males of one shrubland specialist, the Yellow‐breasted Chat (Icteria virens). We banded 1925 juveniles and 2112 adults of nine shrubland species from 2002 to 2005. Of these, 33 (1.7%) juveniles were encountered in subsequent years (2003–2006) as adults (natal dispersal) and 442 (20.9%) birds initially banded as breeding adults were re‐encountered in subsequent years (breeding dispersal). Apparent survival of juvenile shrubland birds on their natal patches was 0.024 (95% CI 0.016–0.036). After accounting for the probability of detection, we found that 21% of birds banded as juveniles and recaptured as adults returned to their natal patches, whereas 78% of adult birds showed fidelity to the patch where they were originally captured. Moreover, natal dispersers tended to move farther than breeding dispersers (corrected natal median = 1.7 km ± 0.37; corrected breeding median = 0.23 km ± 0.10). We used our estimates of natal dispersal and annual apparent survival to estimate true survival at 0.11 (95% CI 0.07–0.18) for juveniles in their first year. However, this estimate was only applicable for birds dispersing within 7 km of their natal patches. Interpatch movements of radio‐marked Yellow‐breasted Chats were not uncommon, with 13 of 37 males located in more than one habitat patch. Overall, we observed low natal philopatry, but high adult site fidelity for shrubland birds in our study area. Considering the frequency of short‐distance movements observed (median = 531 m, range = 88–1045 m), clustering of patches within 1 km might facilitate use of shrubland habitat.     

Fitness consequences of helping behavior in the western bluebird

We examined the fitness consequences of helping behavior inthe western bluebird (Sialia mexicana) at Hastings Reservationin Carmel Valley, California, USA, and tested hypotheses forhow helpers benefit from engaging in alloparental behavior.Both juvenile and adult western bluebirds occasionally helpat the nest During a 12 year period, all adult helpers and mostjuvenile helpers were male. Helpers usually fed at nests ofboth their parents and rarely helped when only one parent waspresent. The frequency of pairs with adult helpers was only7%, but nearly one-third of adult males helped among those withboth parents on the study area. At least 28% were breeders whosenests failed. The propensity to help appears to depend uponparental survival, male philopatry, and the breeding successof potential helpers. Feeding rates were not increased at nestswith juvenile helpers, apparendy because breeding males reducedtheir feeding rates. In contrast, adult helpers increased theoverall rates of food delivery to the nest in spite of a reductionin the number of feeding trips made by both male and femaleparents. Helpers did not derive any obvious direct fitness benefitsfrom helping, but they had greater indirect fitness than nonhelpersdue to increases in nestling growth rates and fledging successat their parents' nests. Helpers fledged fewer offspring intheir first nests than did nonhelpers, suggesting that theywere birds with reduced reproductive potential. Although wehave not yet measured the effect of extrapair fertilizationson the fitness benefits of helping, we calculated the differencein fitness between helpers and nonhelpers as a function of thepotential helper's paternity when breeding independently andhis father's paternity in the nest at which he might help. Inconjunction with constraints on breeding and indirect fitnessbenefits, we predict that relatedness of males to the youngin their own as well as their parents' nests will influencehelping behavior in western bluebirds.     

The Cooperative Breeding System of the Red-cockaded Woodpecker

The breeding system of the red-cockaded woodpecker is described based on data collected over six years from a population of 500 marked individuals in the Sandhills of North Carolina. Male-female pairs were the most common social unit (59%), but 30% of social units contained one or more adult helpers, and 11% consisted of solitary males. Helpers were almost exclusively male: 27% of males remained in their natal group as helpers for at least one year, whereas only four (1%) females did. Most breeding females remained as breeders in the same group from one year to the next (56%), but a surprising number (12%) moved to another group. Many movements were related to incest avoidance or mate death, but 39% involved deserting a mate, usually following successful reproduction. We suggest that females sometimes are forced from groups by immigrants or other group members. The median distance of movements by adult females was only 1.3 km. In contrast to females, no breeding males switched groups. Survival of both breeding (76%) and helper (80%) males was higher than that of breeding females (69%). Males exhibited two distinct life-history strategies. Some remained as helpers on their natal territory for one or more years, and became breeders by inheriting breeding status in the natal group (17% per year) or by replacing a deceased breeder in a nearby group (13% per year, median distance moved 1.0 km). Other males dispersed from their natal group permanently during their first year. Some of these males were floaters at age one year, others were solitary, and a few became helpers in a non-natal group, but many were breeders. In contrast to males that first functioned as helpers, those that dispersed after fledging moved long distances (median dispersal distance 4.5 km), longer even than dispersing female fledglings moved (median distance 3.2 km). The habitat saturation model of the evolution of cooperative breeding is based on selection between the two life-history strategies exhibited by male red-cockaded woodpeckers. The model therefore may be tested directly with this species. Another indication that this model is appropriate for this species is the existence of a resource (cavity trees) that might provide an ecological basis for habitat saturation.     

Reproductive biology of captive male cottontop tamarin monkeys as a function of social environment

The cottontop tamarin, Saguinus oedipus oedipus, is a cooperatively breeding monkey in which mature male and female offspring serve as helpers to assist in rearing younger siblings. Generally, only one female per social group reproduces; breeding restriction is mediated in postpubertal female offspring through low and acyclic levels of reproductive hormones. We investigated (1) reproductive activity of postpubertal male offspring, and (2) whether aggression towards male offspring and a cortisol-mediated stress response might restrict breeding of male offspring in the natal group. We examined sexual behaviour, olfactory communication and urinary hormone levels (testosterone, dihydrotestosterone, luteinizing hormone, cortisol) of the subject males while we manipulated their social environment from housing in natal groups to pairing with a novel female, and after the production of their own offspring. Mounting and erection rates of the male subjects were as high in the natal group as when paired with a novel female. However, most mounts in the natal group were directed towards other males, and complete copulation sequences did not occur with natal-group females. Social environment had no significant effect on olfactory investigation of breeding females. Although hormone levels increased significantly after the subjects were removed from the natal group, the elevation was transient; the hormone levels of subjects in their natal groups did not differ from the levels shown by the same males when successfully producing their own offspring. Male offspring received more contact aggression in the natal group than when paired with the novel female. However, most of the aggression was received from siblings rather than the breeding pair, and levels of cortisol did not correspond with levels of aggression. Thus, at both a behavioural and endocrine level, mature male offspring in captive natal groups were potentially fertile, but sexual activity with natal-group females appeared to be behaviourally restricted and directed instead towards group males. In wild cottontop tamarin groups, this reproductive potential may allow male helpers flexibility to respond to breeding opportunities. Copyright 2001 The Association for the Study of Animal Behaviour.     

Reverse sex-biased philopatry in a cooperative bird: genetic consequences and a social cause

The genetic structure of a group or population of organisms can profoundly influence the potential for inbreeding and, through this, can affect both dispersal strategies and mating systems. We used estimates of genetic relatedness as well as likelihood-based methods to reconstruct social group composition and examine sex biases in dispersal in a Costa Rican population of white-throated magpie-jays ( Calocitta formosa , Swainson 1827), one of the few birds suggested to have female-biased natal philopatry. We found that females within groups were more closely related than males, which is consistent with observational data indicating that males disperse upon maturity, whereas females tend to remain in their natal territories and act as helpers. In addition, males were generally unrelated to one another within groups, suggesting that males do not disperse with or towards relatives. Finally, within social groups, female helpers were less related to male than female breeders, suggesting greater male turnover within groups. This last result indicates that within the natal group, female offspring have more opportunities than males to mate with nonrelatives, which might help to explain the unusual pattern of female-biased philopatry and male-biased dispersal in this system. We suggest that the novel approach adopted here is likely to be particularly useful for short-term studies or those conducted on rare or difficult-to-observe species, as it allows one to establish general patterns of philopatry and genetic structure without the need for long-term monitoring of identifiable individuals.     

Movements of Juvenile Common Ravens in an Arid Landscape

ABSTRACT Movement patterns of juvenile birds are poorly understood, yet critically important ecological phenomena, especially for species with a prolonged juvenile period. We evaluated postfledging movements of juvenile common ravens (Corvus corax) in a western Mojave Desert landscape composed of a mosaic of natural and anthropogenic elements. Generally, ravens do not begin breeding until after their fourth year. We marked 2 annual cohorts of juvenile ravens and followed them from dispersal from their natal territory for up to 33 months. Movements of juvenile common ravens were similar for males and females. Conspecifics and confined livestock feeding operations represented important resources for juvenile ravens, and juveniles were rarely located in open desert. However, initial movements from the natal territory to the nearest communal point subsidy rather than the closest anthropogenic resource suggested juvenile dispersal was influenced by the combination of conspecifics and anthropogenic resources, rather than the distribution of those resources. Land managers concerned with growing raven populations should reduce access to concentrated anthropogenic resources such as landfills and dairies, which serve as important resources for juveniles. Because juvenile ravens rarely venture into open desert, reducing their numbers by lethal removal or other means is unlikely to lessen raven predation of desert tortoises (Gopherus agassizii).     

Departure of juvenile malePresbytis entellus from the natal group

Five bisexual groups (four unimale and one multimale) were studied at Ambagarh Reserve Forest from 1985 to 1987 in relation to factors influencing departures of male juveniles from their natal troops. Of 20 male juveniles observed, 11 left their natal groups, 4 died or disappeared, and 5 remained. The male juveniles left their natal groups only after a takeover by a new adult male. The weaned juvenile males either were chased away by the new adult male or left voluntarily.     

Post-breeding dispersal by female red squirrels (Tamiasciurus hudsonicus): the effect of local vacancies

Red squirrels (Tamiasciurus hudsonicus) defend food-based territoriesyear round, and juveniles must acquire a territory before winterto survive. We experimentally removed territory owners duringthe time that juveniles were becoming independent to examinethe effect of local vacancies on dispersal patterns. Juvenilesattempted to take over removal territories most frequently.However, females with offspring still on the natal territoryactually took over twice as many territories as juveniles. Thesefemales did not appear to move because of low reproductive potentialor to increase territory quality. Instead, moving to a removalterritory allowed more of their offspring to remain on the natalterritory, which appeared to increase juvenile survival.     

Proximate and ultimate causes of natal dispersal in the great bustard Otis tarda

Between 1991 and 1997 we studied the offspring independenceand juvenile dispersal in a wild population of great bustards(Otis tarda). Young males were independent and began their juveniledispersal at an earlier age (6–11 months) than young females(8–15 months). The juvenile dispersal period was longerand the distances reached farther in males than in females.Natal dispersal distances were also longer in males, all ofwhich dispersed from their natal areas and established as adultsat 5–65 km from their natal nests. In contrast, most femaleswere strongly philopatric, settling at 0.5–5 km from theirnatal nests. These marked sex differences in offspring independenceand dispersal may have evolved originally to maintain geneticdiversity and are probably reinforced through male competitionfor mates. Young males that had fed at higher rates and receivedmore feedings from their mothers during the early maternal dependenceperiod became indepthdent and tended to disperse earlier. Theyalso integrated earlier into adult male flocks and settled earlierat their definitive leks, which were closer to their natal sites,in areas of higher adult male density. None of these correlationswas found among young females. These results suggest that enhancedfood intake and maternal care of male offspring are vitallyimportant in increasing their competitive ability during theimmature period and probably also in their fitness as breedingadults. These results are in accordance with the selective valueof large size in males and suggest how this species might havereached such a marked sexual dimorphism in size.     

Social composition of destination territories and matrix habitat affect red-cockaded woodpecker dispersal

We studied the influence of matrix habitats and destination territory social composition on dispersal choices made by red-cockaded woodpeckers (Picoides borealis). We compared matrix habitats and social conditions for 165 dispersals documented between 1995 and 2005 with similarly distanced destinations that went unselected. We modeled data with discrete choice analysis methods, and ranked models using a model selection approach. Model-averaged parameters indicated that juvenile female, and juvenile and helper male red-cockaded woodpeckers dispersed across matrix forests with characteristics similar to those used for breeding. Models differed for sex and social classes, but results generally indicated that birds chose not to transit forests with greater densities of hardwood trees and young pines, and that dispersal was more likely to occur across forests with more large diameter pine trees. Dispersers of both sexes also chose destinations without non-breeding helpers that might present aggressive social challenges to dispersers, particularly males. Juvenile females and helper males also preferred to disperse to territories that were currently occupied by breeding woodpeckers. Previous studies documented the influence on dispersal of travel distance, natal area social interactions and resources, and resources at dispersal destinations. Our results extend factors that affect dispersal movement to include matrix habitats traversed by dispersers and the social composition of destination territories. As such, results have key implications for promoting population connectivity and conservation management of endangered red-cockaded woodpeckers, as well as other resident territorial birds. © 2011 The Wildlife Society.     

Cichlids do not adjust reproductive skew to the availability of independent breeding options

Helpers in cooperatively breeding species forego all or partof their reproduction when remaining at home and assisting breedersto raise offspring. Different models of reproductive skew generatealternative predictions about the share of reproduction unrelatedsubordinates will get depending on the degree of ecologicalconstraints. Concession models predict a larger share when independentbreeding options are good, whereas restraint and tug-of-warmodels predict no effects on reproductive skew. We tested thesepredictions by determining the share of reproduction by unrelatedmale and female helpers in the Lake Tanganyika cichlid Neolamprologuspulcher depending on experimentally manipulated possibilitiesfor helper dispersal and independent breeding and dependingon helper size and sex. We created 32 breeding groups in thelaboratory, consisting of two breeders and two helpers each,where only the helpers had access to a nearby dispersal compartmentwith (treatment) or without (control) breeding substrate, usinga repeated measures design. We determined the paternity andmaternity of 1185 offspring from 47 broods using five to nineDNA microsatellite loci and found that: (1) helpers participatedin reproduction equally across the treatments, (2) large malehelpers were significantly more likely to reproduce than smallhelpers, and (3) male helpers engaged in significantly morereproduction than female helpers. Interestingly, in four broods,extragroup helper males had fertilized part of the brood. Nohelper evictions from the group after helper reproduction wereobserved. Our results suggest that tug-of-war models based oncompetition over reproduction within groups describe best thereproductive skew observed in our study system. Female breedersproduced larger clutches in the treatment compared to the controlsituation when the large helpers were males. This suggests thatmale breeder-male helper reproductive conflicts may be alleviatedby females producing larger clutches with helpers around.     

Male Seychelles warblers use territory budding to maximize lifetime fitness in a saturated environment

In cooperatively breeding species, helping at the nest and buddingoff part of the natal territory have been advanced as strategiesto increase fitness in an environment that is saturated withterritories. The importance of helping or territory buddingas a determinant of lifetime reproductive success (LRS) hasbeen debated because the potential benefits of both strategiescould not be separated. Here we test the causes and the immediateand future fitness consequences of single dispersal decisionstaken by male Seychelles warblers (Acrocephalus sechellensis).Males breeding in high-quality territories (high food abundance)have significantly higher LRS than similar-aged males buddingoff part of the parental territory. Initially, budders havea low reproductive success (because of limited food resourcesor absence of a breeding partner). However, they have a longlife span and inherit high-quality territories through sitedominance, by which they gain higher LRS than breeders on low-qualityterritories, helpers, or floaters. Experimental creation ofmale breeding territory vacancies showed that most young malesbecame budders because of intense competition for high-quality territories. The translocation of warblers to the previouslyunoccupied Aride Island shows that males behave according tothe expected fitness benefits of each dispersal strategy. Inthe absence of competition for territories on Aride, all youngmales bred in high-quality territories. However, after saturationof high-quality habitat with territories, most males becamebudders rather than breeders on low-quality habitat, helpers,or floaters.     

Testis size depends on social status and the presence of male helpers in the cooperatively breeding cichlid Julidochromis ornatus

Cooperatively breeding animals, in which helpers may participatein reproduction with dominant breeders, are ideal species forexamining intraspecific variation in testis size because theyoften exhibit both monogamous breeding (low risk of sperm competition)and polyandrous breeding (high risk) within a population. However,little is known about testis investment as a result of spermcompetition in these animals. The substrate-brooding cichlidfish Julidochromis ornatus has a cooperatively breeding system,in which some males mate monogamously and other males reproduceas dominant breeders or helpers within cooperatively breedinggroups, in which male helpers frequently sire young. We examinedthe relationship between testis investment and male social statusin relation to the risk of sperm competition. As predicted fromsperm competition models, in groups with male helpers, boththe male breeders and the male helpers invested more in testesmass, compared to breeding males without male helpers. We alsofound a positive relationship between the testes mass of malebreeders and their male helpers, suggesting that males increasetheir investment in reproductive capability under the risk ofsperm competition. Sperm competition models also predict thatlarger testes are associated with increased siring success.Our paternity analysis supported this prediction; we found apositive relationship between testis investment by male helpersand the number of offspring they sired.     

Helping by juvenile pine voles (Microtus pinetorum), growth and survival of younger siblings, and the evolution of pine vole sociality

We quantified the behavior of juvenile pine voles (Microtuspinetorum) in families maintained in outdoor enclosures protectedfrom rain and predators and determined the effects of numberof juveniles present (0–3) on their younger siblings andtheir parents. Juveniles exhibited behaviors characteristicof helpers in cooperative breeding species: brooding, grooming,and retrieving younger siblings, and maintaining nests, runways,and food caches. The number of juveniles had no effect on weights,growth rates, or survival of younger siblings or on survivalof parents, but families with three juveniles had shorter interlitterintervals than other families. There was a trend toward lowerweights for younger siblings when juveniles continued to nursefrom their mothers. Juveniles brooded and groomed younger siblingsmore when their mothers were absent than when mothers were present.In families with juveniles, younger siblings were alone significandyless often than in families without juveniles, but number ofjuveniles in the family had little effect on duration or frequencywith which juveniles brooded or groomed younger siblings. Thesedata are consistent with cooperative breeding having evolvedvia individual selection for delayed dispersal. We argue thatthe limited availability of vacant tunnel systems and the highcost of digging new tunnel systems constrains dispersal in pinevoles and has led to the evolution of cooperative breeding.     

Delayed juvenile dispersal benefits both mother and offspring in the cooperative spider Anelosimus studiosus (Araneae: Theridiidae)

Anelosimus studiosus juveniles usually remain in their natalwebs with their mothers until maturity, forming temporary coloniesin which individuals cooperate in web maintenance and preycapture. In a semi-natural environment, we experimentally removedjuveniles from their natal webs at mid-development. In thecontrol group, the juveniles were immediately replaced in theirnatal webs; in the experimental removal group, the juvenileswere not replaced and a sample of them were allowed to buildindividual webs. Colonies and solitary juveniles were exposedto natural prey densities and censused regularly for numbersand stages of spiders, and for prey capture. On average, juvenilesin colonies survived longer, developed faster, and had more resources per individual than did solitary juveniles. However,some of the solitary juveniles obtained more resources thanindividual juveniles in colonies. Mothers in the control groupsurvived longer and produced second broods earlier than mothersin the experimental removal group. Within the control group,older and larger colonies captured more and larger prey. Larger colonies had a lower coefficient of variation in prey capturedper juvenile. Overall, delayed juvenile dispersal benefitsboth juveniles and mothers.     

Sex-bias and timing of natal dispersal in cooperatively breeding Puff-throated Bulbuls Alophoixus pallidus

While natal dispersal can have a significant impact on population dynamics, it is typically difficult to quantify. We investigated timing of natal dispersal of the cooperatively breeding Puff-throated Bulbul Alophoixus pallidus in a tropical evergreen forest by modelling the probability of staying in or dispersing from their natal territory whilst taking into account the effects of sex, group size, and the presence of helper(s). Birds did not disperse until the beginning of and during the breeding season following the hatching year. Dispersal was strongly female-biased both in frequency and distance: most females (95%) dispersed away from their natal territories, and of those relocated, traversed 2–7 territories. In contrast, 50% of males remained in the natal territory as helpers in their second year, while relocated dispersing males crossed 1–2 territories. Natal dispersal was not influenced by either group size or the presence of helpers. Males that fledged earlier in the breeding season exhibited higher rates of philopatry than the males that fledged later, but no correlation between fledging date and philopatry was observed in females. The probability of staying in the natal territory during the second year was 0.58 ± 0.14 SE and 0.05 ± 0.04 for males and females, respectively. These findings may add to our understanding of how natal dispersal can reflect social patterns and kin structure in cooperative breeding species from a little-studied tropical forest region.     

Paying to stay or paying to breed? Field evidence for direct benefits of helping behavior in a cooperatively breeding fish

Several hypotheses aim to explain the evolution of helping behavior,but conclusive experimental support for evaluating the relativeimportance of individual hypotheses is still lacking. We reporton two field experiments conducted to test the "territory inheritance"and "pay-to-stay" hypotheses in the cooperatively breeding cichlidfish Neolamprologus pulcher The territory inheritance hypothesiswas tested by removing one parent, which created breeding vacancies.In 39% of cases, same-sex helpers took over the breeding spot;in 44% of cases helpers continued helping new breeders, and17% were evicted by new breeders. Helpers that were closelysize matched to the removed breeder had a better chance of gainingthe breeding spot Male helpers tended to continue helping aftera takeover more often than females.The pay-to-stay hypothesiswas tested-by temporarily removing helpers. Whereas breedersdid not respond aggressively to removals, other group membersattacked the removed helpers on their return, and 29% were eventuallyevicted. The returning helpers assisted more by increasing theirrate of territory maintenance and defense and visiting the broodchamber more frequently Size and sex of removed helpers didnot explain the observed aggressive reactions of other groupmembers. Thus, our results support both hypotheses: N. pulcherneeds to pay with help to be allowed to remain protected inthe family group, and there they may inherit the natal territory.N. pulcher helpers gain direct benefits from helping behavior.     

Survival of Juvenile Pygmy Rabbits

ABSTRACT Until recently, natal behavior of pygmy rabbits (Brachylagus idahoensis) was largely unknown, and no information on survival of free-ranging juveniles was available. We evaluated survival of radiotagged juvenile pygmy rabbits at 2 sites in east-central Idaho, USA, during 2004 and 2005. We captured juveniles (n 58) shortly after they emerged from natal burrows. Mortality rates were high and variable, ranging from 27% for females during 2004 to 63% for males during 2005. Approximately 69% of mortalities were attributed to predation. We evaluated variables influencing juvenile survival through 18 weeks old using known-fate models in Program MARK. We expected survival to decline around the age of natal dispersal and to be lower for young born later in the season. We evaluated 14 candidate models that included sex, year, study area, and relative date of birth within each year. Model selection results did not indicate strong support for any single combination of variables, and 8 competing models all included effects of relative date of birth, year, and study area. These results revealed substantial variability in survival of juveniles across multiple factors, and we documented similar patterns for adult pygmy rabbits. Such high variability in survival over relatively small spatial and temporal scales might contribute to marked fluctuations in populations of pygmy rabbits and, hence, managers interested in monitoring this species might consider monitoring multiple populations across broader geographic areas to assess regional trends in numbers.     

Malformed offspring,sibling matings,and selection against inbreeding in the sand lizard (Lacerta agilis)

We demonstrated that sand lizards (Lacerta agilis) are more likely to have malformed offspring when they mate with siblings. Offspring with malformations, such as deformed limbs and heads, have zero survival in a natural population. Normal-looking siblings of malformed hatchlings also had a reduced survival in the wild, compared to offspring from clutches in which all siblings appeared normal. The proportion of malformed hatchlings in the natural population was ca. 10%, in spite of differences in juvenile dispersal between males and females. Male juveniles disperse significantly further from their natal sites than do female juveniles.     

Spatial genetic relationships in a population of the Japanese wood mouse Apodemus argenteus

Space-related relationships between individuals were investigated during the breeding season in a natural population of Apodemus argenteus (Temminck) by using the catch-mark-release method and microsatellite DNA markers. Several overlapping patterns of home ranges in breeding males and females were observed. Although this species is thought to be monogamous, DNA analysis revealed polygynous mating. In adults, relatedness between sexes was generally low and no apparent spatial genetic structure was observed. In juveniles, however, relatedness and geographical distance be-tween individuals were significantly negatively correlated. The relatedness between adult females decreased with distance, whereas there was not such a correlation in males. Long-distance movements were observed in current-year-born males. The differences in spatial genetic structure between adults and juveniles are thought to be produced by male-biased natal dispersal. An erratum to this article can be found at