Postfledging habitat selection of juvenile middle spotted woodpeckers: a multi-scale approach

Despite its relevance for the persistence of populations, the ecological mechanisms underlying habitat use decisions of juvenile birds are poorly understood. We examined postfledging habitat selection of radio-tracked juvenile middle spotted woodpeckers Dendrocopos medius at multiple hierarchically-nested spatial scales in NW Spain. At the landscape and home range scales, old oak forest was the most used and selected habitat, young oak forests and pine plantations were avoided, and riverside forests were used as available. At a lower scale, birds selected larger diameter trees for foraging. Home ranges had higher densities of large deciduous trees (mainly oaks Quercus spp., but also poplars Populus spp. and willows Salix spp. >22  cm and >33  cm DBH) selected for foraging by juveniles than non-used areas. These results suggest that foraging conditions may drive, at least partly, habitat use decisions by juvenile birds. We also discuss the potential influence of intraspecific competition, the search for a future breeding territory in the early postfledging period and predation avoidance on habitat use decisions by juvenile birds. Contrary to previous studies on migrant forest birds, postfledging juvenile woodpeckers selected the same habitat as for the breeding adults (i.e. old oak forest), indicating that migrant and resident specialist avian species may require different conservation actions. Conservation strategies of woodpecker populations should consider the protection of old oak forests with high densities of large trees to provide suitable habitat to breeding adults and postfledging juveniles. The habitat improvement for this indicator and umbrella species would also favour other organisms that depend on characteristics of old-growth oak forests.     

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.     

Heart rot hotel: fungal communities in red-cockaded woodpecker excavations

Tree-cavity excavators such as woodpeckers are ecosystem engineers that have potentially complex but poorly documented associations with wood decay fungi. Fungi facilitate cavity excavation by preparing and modifying excavation sites for cavity excavators. Associations between fungi and endangered red-cockaded woodpeckers (RCWs) are particularly interesting because these are the only birds that specialize in excavating into the heartwood of living pines, a process that takes years to complete. Using molecular methods, we examined fungal communities in complete and incomplete RCW excavations, and non-cavity control trees. In addition to finding a high diversity of fungi, we found three groupings of fungal communities corresponding to the three groups of trees sampled. We show that trees selected for cavity excavation by RCWs are infected by distinct fungal communities, and propose two hypotheses to explain this outcome: the bird facilitation hypothesis and the tree selection hypothesis.     

The breeding ecology and behaviour of Wrens Troglodytes troglodytes under primaeval and secondary conditions

The breeding habits of Wrens inhabiting the primaeval forest of Biatowieza National Park, Poland, are described and compared with those of Wrens living in man-transformed habitats of Western Europe.
Significant inter-habitat differences in numbers exist in Biatowieza. On average there are 4-3 territories per 10 ha in ash-alder, 1 -7 territories per 10 ha in oak–hornbeam and 0–4 territories per 10 ha in coniferous habitats. In ash–alder stands the males have smaller territories, build fewer nests and have better pairing success than in oak–hornbeam stands. However, these two habitats do not differ significantly in average clutch size, timing of breeding or nesting success. Over 80% of nests used for breeding in Biatowieza are situated in uprooted trees. Inter-habitat differences in bird numbers reflect differences in distribution and density of such trees.
Low numbers of Biatowieza Wrens cannot be accounted for solely by high mortality away from the breeding grounds. Available data strongly suggest that numbers of both sexes are proximately limited by behavioural interactions on the breeding ground.
In west European Wrens, densities are higher, territories smaller, polygyny more frequent, clutches smaller and production of young higher than in Biatowieza. Differences in production of young result from different rates of nest predation. Indirect evidence suggests that the breeding habits of Wrens once living in the primaeval forests of western Europe were similar to those of Biatowieza birds, and that the differences observed nowadays result to a large extent from man-made transformations of habitat. It is suggested that man's lowering of the predation pressure acting both on proximate (enabling birds to produce more young) and ultimate levels (decreasing relative advantages of dispersed breeding) was the decisive factor allowing an increase in numbers of Wrens in secondary habitats.     

Stepping stones of life: natal dispersal in the group-living but noncooperative speckled warbler

In most cooperatively breeding birds the offspring of one sex, usually male, delays dispersal to remain on the natal territory and helps its parents to rear subsequent young. Thus delayed dispersal could be the first step in the evolution of cooperative breeding. We studied natal dispersal in a population of the group-living speckled warbler, Chthonicola sagittata, based on observations of a colour-banded population over 3 years. Unlike other group-living members of the Acanthizinae, all juvenile males in this population dispersed to settle on foreign territories as subordinates, which do not help rear the young. Speckled warblers showed all the life history traits that are thought to result in a saturated habitat and lead to delayed dispersal: they were sedentary, had high adult survival and had a male-biased sex ratio. However, they differed from other acanthizids in occurring at low density (0.18 birds/ha) on large breeding territories (6-12 ha), with a maximum of two males per territory. This may allow subordinates to live on foreign territories yet avoid aggression from dominants. A benefit of dispersal is that it provides an additional route to gaining a breeding vacancy. Dispersers can acquire vacancies on their new territory or on a neighbour's, but incest avoidance would be likely to constrain nondispersing males to neighbours' territories. A model of relative lifetime success showed that the survival benefits of natal philopatry are unlikely to outweigh this benefit of dispersal.     

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.     

Seed Dispersal by Birds and Bats in Lowland Philippine Forest Successional Area

In the tropical forests of SE Asia, only a few studies have dealt with the role animal dispersal plays in early forest succession and rehabilitation, and a comparison of bird and bat dispersal is even rarer. We investigated seed dispersal by birds and bats in a successional area in the lowland dipterocarp forest of the Subic Watershed Forest Reserve (SWFR) in Luzon Island, Philippines. Using pairs of day and night traps, we collected seeds during 3 mo of wet season and 3 mo of dry season in a 1.2-ha study site. Bird-dispersed seeds predominated over those dispersed by bats in terms of both seed abundance and number of seed species. The most abundant endozoochorous seed species were significantly biased toward either bird or bat dispersal. Birds and bats appeared to compete more strongly for fruit resources during the dry season than during the wet season, and bats responded more to changes in the seasons than birds did. GLM analyses showed that the factor that had the strongest influence on overall seed distribution was the number of fleshy-fruited trees surrounding the traps, and that the distribution pattern of day-dispersed seeds was affected by more physical factors (number of trees, size of trees, presence of fleshy-fruited and conspecific trees) in the study site than the pattern of night-dispersed seeds were. Given that birds are the more important dispersers in the study site, restoration efforts in SWFR might benefit by focusing on attracting these dispersers into its degraded habitats.     

High-quality seed dispersal by fruit-eating fishes in Amazonian floodplain habitats

Seed dispersal is a critical stage in the life history of plants. It determines the initial pattern of juvenile distribution, and can influence community dynamics and the evolutionary trajectories of individual species. Vertebrate frugivores are the primary vector of seed dispersal in tropical forests; however, most studies of seed dispersal focus on birds, bats and monkeys. Nevertheless, South America harbors at least 200 species of frugivorous fishes, which move into temporarily flooded habitats during lengthy flood seasons and consume fruits that fall into the water; and yet, we know remarkably little about the quality of seed dispersal they effect. We investigated the seed dispersal activities of two species of large-bodied, commercially important fishes (Colossoma macropomum and Piaractus brachypomus, Characidae) over 3 years in Pacaya-Samiria National Reserve (Peru). We assessed the diet of these fishes during the flood season, conducted germination trials with seeds collected from digestive tracts, and quantified fruit availability. In the laboratory, we fed fruits to captive Colossoma, quantified the proportion of seeds defecated by adult and juvenile fish, and used these seeds in additional germination experiments. Our results indicate that Colossoma and Piaractus disperse large quantities of seeds from up to 35% of the trees and lianas that fruit during the flood season. Additionally, these seeds can germinate after floodwaters recede. Overexploitation has reduced the abundance of our focal fish species, as well as changed the age structure of populations. Moreover, older fish are more effective seed dispersers than smaller, juvenile fish. Overfishing, therefore, likely selects for the poorest seed dispersers, thus disrupting an ancient interaction between seeds and their dispersal agents. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Frugivory,Post‐feeding Flights of Frugivorous Birds and the Movement of Seeds in a Brazilian Fragmented Landscape

Habitat fragmentation can break down the movement processes of frugivorous animals, thus influencing the relationship between plants and their seed dispersers by altering the number and identity of seed dispersers, and their relative contribution to seed dispersal. We studied the assemblages of frugivorous birds, their composition, species richness, and visitation rates to fruiting plants growing in the different landscape elements (forest fragments, live fences, and trees isolated in pastures) embedded in a Brazilian fragmented, agricultural landscape. By following the post‐feeding movements of frugivorous birds, we inferred the direction of seed movement from and to each of these landscape elements. Fruiting trees growing at different landscape elements were visited by frugivorous birds at similar rates. Isolated trees attracted a greater and distinct bird assemblage than trees in forest fragments or live fences. Judging by the post‐feeding flights of birds, the seeds of isolated trees were the most likely to reach all the landscape elements considered, but the contribution of isolated trees to the seeds falling in forested habitats or pastures depended on their degree of isolation. A few bird species were able to move widely, visiting fruiting plants in all landscape elements, and promoting long‐distance dispersal for plants. These few birds are of special interest because they are mobile links that connect habitats in fragmented landscapes with their seed dispersal services. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

Body size and social dominance influence breeding dispersal in male Pachydiplax longipennis (Odonata)

1. Dispersal behaviour can be affected by an individual's phenotype, by the environmental or social context they experience, and by interactions between these factors. Differential dispersal propensities between individuals may also be an important modifier of functional connectivity between populations. To assess how a key trait, body size, affected both social interactions and dispersal behaviour, this study examined the relationship between body size, antagonistic interactions, and breeding dispersal in male dragonflies (Pachydiplax longipennis) across a seasonal decline in adult body size. 2. During a seasonal peak in male body size in this study, dispersers were smaller than non‐dispersers. Later in the season, the body size of dispersers and non‐dispersers did not differ. 3. Focal observations found that body size was related to competitive dominance, large males engaged in aggressive chases more often and smaller males were more frequently pursued. 4. These results indicate that when large males were present, small males were more likely to disperse suggesting that dispersal is a tactic adopted by social subordinates in this context. If breeding dispersal is typically undertaken by subordinate males, functional connectivity between populations may be less than estimated from absolute dispersal rates.     

Natal dispersal in great bustards: the effect of sex, local population size and spatial isolation

1. We investigated the causes of natal dispersal in four Spanish areas where 35 breeding groups of the polygynous great bustard Otis tarda were monitored intensively. A total of 392 juveniles were radio-tracked between 1991 and 2006 by ground and via aeroplane to avoid potential biases derived from the non-detection of long-distance dispersers. 2. We explored 10 explanatory variables that were related to individual phenotypic features, habitat and conspecific traits in terms of group size and breeding performance, and spatial distribution of available breeding groups. Probability of group change and natal dispersal distances were investigated separately through multifactorial analyses. 3. Natal dispersal occurred in 47.8% of the birds and median natal dispersal distance of dispersers was 18.1 km (range 4.97-178.42 km). Sex largely determined the dispersal probability, with 75.6% of males being dispersers and 80.0% of females being philopatric, in contrast to the general pattern of female-biased dispersal found in most avian species. 4. Both the frequency of natal dispersal and dispersal distances were affected by the spatial distribution of breeding groups. More isolated groups showed a higher proportion of philopatric individuals, the effect being more evident in males than in females. This implies a reduction in gene flow in fragmented populations, as most genetic exchange is achieved through male dispersal. Additionally, dispersers hatched in more isolated groups tended to exhibit longer dispersal distances, which increases the associated energetic costs and mortality risks. 5. The dispersal decision was influenced by the number of conspecifics in the natal group. The individual probability of natal dispersal was related inversely to the size of the natal group, which supports the balanced dispersal model and the conspecific attraction hypothesis. 6. Overall, our results provide a good example of phenotypic plasticity and reinforce the current view that dispersal is an evolutionary complex trait conditioned by the interaction of individual, social and environmental causes that vary between individuals and populations.     

Habitat exploration and use in dispersing juvenile flying squirrels

1. Variation in behaviours involved in habitat selection is important for several evolutionary and ecological processes. For example, habitat use during dispersal may differ from breeding habitat use, and for dispersers the scale of habitat familiarity is determined by exploratory behaviour. We studied habitat use and exploration of 56 radio-collared juvenile flying squirrels Pteromys volans L. within natal home range and during dispersal, and compared habitat use between juveniles and 37 adults within breeding home range. 2. Before dispersal, young flying squirrels actively moved around the natal site. Surprisingly, long-distance dispersers explored less than short-distance dispersers, but philopatric individuals explored similar distances as dispersers. Females explored less than males, although females are the more dispersive sex in flying squirrels. 3. For most of the individuals the settlement area was unfamiliar due to long dispersal distance. Consequently, direction and distance of exploration were not very strong predictors of settlement location. However, individuals familiar with the settlement area concentrated exploration to that area. Exploration did not correlate with short-term survival. 4. Dispersers preferred breeding habitat while dispersing, but were found more often in matrix habitat than juveniles within natal, or adults within breeding, home ranges. 5. We conclude that familiarity does not determine settlement as much as, for example, availability of the habitat for flying squirrels. Based on our results, it also seems clear that data on adult habitat use are not enough to predict habitat use of dispersing individuals. In addition, our results support the recent view that short- and long-distance dispersers may need to be analysed separately in ecological and evolutionary analyses.     

Genetic monogamy in single-male groups of acorn woodpeckers, Melanerpes formicivorus

Acorn woodpeckers have one of the most complex social systems of any bird species. Breeding units range in size from monogamous pairs to groups of 15 birds that include multiple breeding males and females as well as nonreproductive helpers-at-the-nest. Groups form when young remain at their natal nest to help their parents breed or when single-sex coalitions of siblings disperse to fill a reproductive vacancy on another territory. Plural breeding and helping behaviour are thought to be favoured through indirect fitness benefits for individuals that would otherwise be unable to breed due to a shortage of reproductive vacancies on territories with acorn stores. We report the results of multi-locus DNA fingerprinting of 51 offspring from 18 nests of 16 socially monogamous pairs of acorn woodpeckers. If socially monogamous females mate outside the pair-bond, indirect fitness benefits for cobreeders and helpers will be significantly reduced. Monogamous pairs accounted for all but one of the 51 offspring we tested; the single exception was apparently sired by the putative father, but the putative mother was excluded from maternity. Our results indicate that individuals remaining on their natal territories as helpers are generally the genetic offspring of the pair they help. They also suggest that single-sex coalitions offspring dispersing together from nests of socially monogamous pairs will be full-siblings.     

Competition for mates and predominant juvenile male dispersal in mammals

Competition among males for mates may partially explain the predominant dispersal of juvenile male California ground squirrels (Spermophilus beecheyi). I tested the generality of this hypothesis by comparing dispersal patterns in many species of mammals. In most polygynous and promiscuous species of mammals, juvenile males are the predominant dispersers. In most monogamous species of mammals, juvenile males and juvenile females are predominant dispersers with little distinction as to sex. While many factors may influence dispersal, achieving outcrossing and advantages in competition for mates are most likely to explain predominant dispersal by juvenile males in many mammal species.     

千岛湖岛屿社鼠的扩散特征

2009年7月—11月,2010年3月—11月对千岛湖两个岛屿上的社鼠(Niviventer confucianus)种群进行标志重捕,通过分析扩散个体的性别、年龄及扩散距离等,探讨陆桥岛屿环境下社鼠的扩散特征。研究结果显示:岛屿环境下社鼠的扩散率为37.96%,不同季节社鼠的扩散率存在差异,社鼠倾向于在春季发生扩散,尤其是3月份,社鼠扩散个体数最多;对于整个捕获史而言,雌雄扩散个体数存在极显著的差异(x~2=12.542,df=1,P0.001),雄性个体更倾向于发生扩散;而在繁殖高峰期,当怀孕鼠存在时,雄性扩散数亦大于雌性,两者存在显著差异(x~2=4.504,df=1,P=0.034)。扩散个体体重的季节差异不显著;扩散与社鼠的年龄密切相关,社鼠更加倾向于在亚成年阶段发生扩散;另外,家群大小对社鼠扩散个体数量也有显著影响,家群越大,扩散个体越多;对扩散距离进行分析发现存在显著的性别差异(t=-9.229,df=50,P0.001),雌性个体的平均扩散距离为(282.97±16.14)m,显著大于雄性的(146.70±6.80)m,而扩散距离与年龄、体重、最大活动距离均无显著相关性。     

EXTRAPAIR MATE CHOICE AND HONEST SIGNALING IN COOPERATIVELY BREEDING SUPERB FAIRY-WRENS

In many species of monogamous birds females copulate with males other than their social mates, resulting in extrapair fertilizations. Little is known about how females choose extrapair mates and whether the traits used to choose them are reliable indicators of male quality. Here we identify a novel male trait associated with extra-group mating success in the superb fairy-wren (Malurus cyaneus), a cooperatively breeding bird with one of the highest known frequencies of extra-group mating. Female fairy-wrens chose extra-group mates that molted earlier into breeding plumage. Males molted up to five months before the breeding season began, and only males that molted at least one month prior to its onset gained any extra-group fertilizations. This conclusion held after controlling statistically for the effect of age and social status on molt date. Once males acquired breeding plumage, they began courtship display to females on other territories. Thus, some males were displaying to females for several months before the breeding season began. This extraordinarily long period of advertisement by males may be facilitated by the long-term ownership of territories. We suggest that early acquisition of breeding plumage or the subsequent display behavior can be reliable cues for mate choice because they are costly to acquire or maintain.     

Plumage brightness predicts non-breeding season territory quality in a long-distance migratory songbird, the American redstart Setophaga ruticilla

Many species of birds exhibit brilliant ornamental plumage, yet most research on the function and evolution of plumage has been confined to the breeding season. In the American redstart Setophaga ruticilla , a long-distance Neotropical-Nearctic migratory bird, the acquisition of a winter territory in high-quality habitat advances spring departure and subsequent arrival on breeding areas, and increases reproductive success and annual survival. Here, we show that males holding winter territories in high-quality, black mangrove habitats in Jamaica have brighter yellow-orange tail feathers than males occupying territories in poor-quality second-growth scrub habitats. Moreover, males arriving on the breeding grounds from higher-quality winter habitats (inferred by stable-carbon isotopes) also had brighter tail feathers. Because behavioral dominance plays an important role in the acquisition of winter territories, plumage brightness may also be related to fighting ability and the acquisition and maintenance of territories in high-quality habitat. These results highlight the need for further research on the relationships between plumage coloration, behavior, and the ecology of over-wintering migratory birds.     

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.     

Dispersal modes affect tropical forest assembly across trophic levels

We examined assemblages of trees and two major groups of vertebrate seed dispersers, birds and primates, in Ugandan protected areas to evaluate the roles of dispersal limitation and species sorting in community assembly. We conducted partial Mantel tests to investigate relationships between community similarity, environmental distance and geographic distance. Results showed that environmental factors, specifically temperature and rainfall, significantly and more strongly structured tree assemblages than geographic distance. Analysis of tree dispersal modes revealed wind‐dispersed tree guilds were significantly dispersal limited but trees dispersed by animals were not. For assemblages of vertebrate seed dispersers, dispersal limitation significantly and more strongly structured assemblages of primates than species sorting whereas environmental factors significantly and more strongly structured assemblages of birds than dispersal limitation. We therefore examined whether trees dispersed by primates were more dispersal limited than trees dispersed by birds. We found consistent trends that primate fruit trees were more dispersal limited than bird fruit trees using three definitions of dispersal syndromes based on fruit color. Our results suggest that the dispersal abilities of primary consumers may affect the distribution of primary producers at large spatial scales.