Social Behavior,Reproductive Strategies,and Population Genetic Structure of <Emphasis Type="Italic">Lagothrix poeppigii</Emphasis>

For species of primates in which females emigrate, we would expect males within groups to be related to one another. Kin selection theory suggests that these males should associate preferentially with one another, be more affiliative and cooperative with one another than females are, and compete less overtly with one another over reproductive opportunities than males in female philopatric taxa do. Precisely these patterns of social behavior characterize well-studied populations of 2 of the 3 atelin primate genera: spider monkeys (Ateles) and muriquis (Brachyteles). For the third atelin genus, Lagothrix, patterns of intragroup social behavior have been less well-documented. We studied the social and reproductive behavior of lowland woolly monkeys (Lagothrix lagotricha poeppigii) in Ecuador during a one-year observational study and subsequently used molecular techniques to investigate population genetic structure and dispersal patterns for this taxon. Among adult male woolly monkeys, both affiliative and agonistic interactions were rare, and males were seldom in close proximity to one another. Relationships among male woolly monkeys are best characterized as tolerant, especially in the context of mating wherein direct competition among males was minimal despite the fact that females mated with multiple males. Relationships among females were likewise generally tolerant but nonaffiliative, though females often directed harassment towards copulating pairs. Affiliative interactions that did occur among woolly monkeys tended to be directed either between the sexes—primarily from female to male—or from younger towards older males, and the proximity partners of females tended to be members of the opposite sex. These results suggest that bonds between the sexes may be more important than same-sex social relationships and that direct female-female competition is an important feature of woolly monkey reproductive biology. Our genetic results indicate that, as in other atelins, dispersal by females is common, but some male dispersal likely occurs as well. In some but not all groups we studied, nonjuvenile males within social groups were more closely related to one another on average than females were, which is consistent with greater male than female philopatry. However, differences in these patterns among our study groups may reflect local variation in dispersal behavior.     

Social constraint and an absence of sex‐biased dispersal drive fine‐scale genetic structure in white‐winged choughs

This study used eight polymorphic microsatellite loci to examine the relative effects of social organization and dispersal on fine‐scale genetic structure in an obligately cooperative breeding bird, the white‐winged chough (Corcorax melanorhamphos). Using both individual‐level and population‐level analyses, it was found that the majority of chough groups consisted of close relatives and there was significant differentiation among groups (F_ST = 0.124). However, spatial autocorrelation analysis revealed strong spatial genetic structure among groups up to 2 km apart, indicating above average relatedness among neighbours. Multiple analyses showed a unique lack of sex‐biased dispersal. As such, choughs may offer a model species for the study of the evolution of sex‐biased dispersal in cooperatively breeding birds. These findings suggest that genetic structure in white‐winged choughs reflects the interplay between social barriers to dispersal resulting in large family groups that can remain stable over long periods of times, and short dispersal distances which lead to above average relatedness among neighbouring groups.     

Dictyostelids from Mediterranean forests of the south of Europe

The first results of a study of dictyostelid diversity in soils of Mediterranean vegetation are presented. Surface soil samples were collected during 2003–2004 from different Mediterranean forests in Spain and Portugal and were plated out for cellular slime moulds. Isolated during this study were 14 species of dictyostelids. One species, Dictyostelium firmibasis is reported for the first time from Europe, and nine (Dictyostelium aureo-stipes, Dictyostelium sp. 1, Dictyostelium fasciculatum, Dictyostelium giganteum, Dictyostelium implicatum, Dictyostelium leptosomum, Dictyostelium sphaerocephalum, Polysphondylium candidum, Polysphondylium pallidum) are new records for the Mediterranean region. Members of the three genera Dictyostelium (ten species), Polysphondylium (three species) and Acytostelium (one species) are present in this region of the world. Comments on their diversity and distribution are included.     

Phylogeography and sexual macrocyst formation in the social amoeba <Emphasis Type="Italic">Dictyostelium giganteum</Emphasis>

Background  

Microorganisms are ubiquitous, yet we are only beginning to understand their diversity and population structure. Social amoebae (Dictyostelia) are a diverse group of unicellular eukaryotic microbes that display a unique social behaviour upon starvation in which cells congregate and then some die to help others survive and disperse. The genetic relationships among co-occurring cells have a major influence on the evolution of social traits and recent population genetic analysis found extensive genetic variation and possible cryptic speciation in one dictyostelid species (Dictyostelium purpureum). To further characterize the interplay among genetic variation, species boundaries, social behaviour, and reproductive isolation in the Dictyostelia, we conducted phylogenetic analyses and mating experiments with the geographically widespread social amoeba Dictyostelium giganteum.     

When dispersal fails: unexpected genetic separation in Gibraltar macaques (Macaca sylvanus)

Barbary macaques (Macaca sylvanus), now restricted in the wild to a few isolated forested areas of Morocco and Algeria, are present in a free‐ranging colony on Gibraltar. For many decades, the Gibraltar colony was exposed to multiple bottlenecks due to highly nonrandom removal of animals, followed by repeated introductions of animals from North Africa. Moreover, because of complete isolation, Gibraltar's several social groups of macaques provide an ideal system to study the genetic consequences of dispersal in cercopithecines in situ. Predictions of genetic consequences due to male‐biased dispersal in cercopithecines will be different for autosomal and maternally inherited genetic markers, such as the control region of the mitochondrial DNA. We used a panel of 14 highly polymorphic microsatellite loci and part of the hypervariable region I of the mitochondrial control region to estimate genetic structure between five social groups in Gibraltar. Surprisingly, for autosomal markers, both classical summary statistics and an individual‐based method using a Bayesian framework detected significant genetic structure between social groups in Gibraltar, despite much closer proximity than wild Algerian and Moroccan populations. Mitochondrial data support this finding, as a very substantial portion of the total genetic variation (70.2%) was found between social groups. Using two Bayesian approaches, we likewise identified not only a small number of male first‐generation immigrants (albeit less than expected for cercopithecines) but also unexpectedly a few females. We hypothesize that the culling of males that are more likely to disperse might slow down genetic homogenization among neighbouring groups, but may also and more perversely produce selection on certain behavioural traits. This may have important repercussions for conservation, as it could lead to evolutionary changes that are not due to inbreeding or genetic drift.     

Dispersal patterns in black howler monkeys (Alouatta pigra): Integrating multiyear demographic and molecular data

Dispersal is a fundamental process in the functioning of animal societies as it regulates the degree to which closely related individuals are spatially concentrated. A species’ dispersal pattern can be complex as it emerges from individuals’ decisions shaped by the cost–benefit tradeoffs associated with either remaining in the natal group or dispersing. Given the potential complexity, combining long-term demographic information with molecular data can provide important insights into dispersal patterns of a species. Based on a 15-year study that integrates multiyear demographic data on six groups with longitudinal and cross-sectional genetic sampling of 20 groups (N = 169 individuals, N = 21 polymorphic microsatellite loci), we describe the various dispersal strategies of male and female black howler monkeys (Alouatta pigra) inhabiting Palenque National Park, Mexico. Genetically confirmed dispersal events (N = 21 of 59 males; N = 6 of 65 females) together with spatial autocorrelation analyses revealed that the dispersal pattern of black howlers is bisexual with strong sex-biases in both dispersal rate (males disperse more often than females) and dispersal distance (females disperse farther than males). Observational and genetic data confirm that both males and females can successfully immigrate into established groups, as well as form new groups with other dispersing individuals. Additionally, both males and females may disperse singly, as well as in pairs, and both may also disperse secondarily. Overall, our findings suggest multiple dispersal trajectories for black howler males and females, and longer multiyear studies are needed to unravel which demographic, ecological and social factors underlie individuals’ decisions about whether to disperse and which dispersal options to take.     

Fine-scale analysis of genetic structure in the brooding coral Seriatopora hystrix from the Red Sea

The dispersal of gametes and larvae plays a key role in the population dynamics of sessile marine invertebrates. Species with internal fertilisation are often associated with very localised larval dispersal, which may cause small-scale patterns of neutral genetic variation. This study on the brooding coral Seriatopora hystrix from the Red Sea focused on the smallest possible scale: Two S. hystrix stands (~100 colonies each) near Dahab were completely sampled, mapped and analysed at five microsatellite markers. The sexual mode of reproduction, the likely occurrence of selfing and the level of immigration were in agreement with previous studies on this species. Contrary to previous findings, both stands were in Hardy–Weinberg proportions. Also, no evidence for spatially restricted larval dispersal within the sampled areas was found. Differences between this and previous studies on S. hystrix could reflect variation in life history or varying environmental conditions, which opens intriguing questions for future research.     

Fine‐scale genetic structuring in a group‐living lizard,the gidgee skink (Egernia stokesii)

By constraining gene flow, group living and natal philopatry can result in fine‐scale genetic structure. Although the genetic structure of some group‐living lizards has been characterised, studies are few compared with those for group‐living bird and mammal species. The Egerniinae group of lizards exhibits a high diversity of social structures, making it a useful group for comparative studies of genetic structure across a broader range of social taxa. A well‐studied member of Egerniinae is Egernia stokesii, a lizard that forms long‐term pair bonds and stable social groups and exhibits natal philopatry and limited dispersal. Evidence exists for consistent E. stokesii social structure across seven close but disconnected rocky outcrops within a 40 × 10 km area. We used summary statistics, analysis of molecular variance, Bayesian clustering, and discriminant analysis of principal components to assess if E. stokesii exhibit a consistent pattern of fine‐scale genetic structure across the same seven outcrops. Due to E. stokesii social structure and constrained dispersal, we predicted significant genetic structuring – based on microsatellite markers – among outcrops. We found significant fine‐scale genetic structuring and evidence for two genetic clusters. We discuss features of E. stokesii biology and ecology that may explain our findings. Some rocky outcrops, and some social groups, contained lizards from both genetic clusters. An examination of the composition of mixed cluster social groups did not detect any notable patterns. Therefore, further work is necessary to identify how the observed patterns may have arisen. Future investigations in E. stokesii and other group‐living lizard species are likely to contribute greatly to our understanding of the genetic consequences of group living.     

Research progress in avian dispersal behavior

Dispersal, defined as a linear spreading movement of individuals away from others of the population is a fundamental characteristic of organisms in nature. Dispersal is a central concept in ecological, behavioral and evolutionary studies, driven by different forces such as avoidance of inbreeding depression, density-dependent competition and the need to change breeding locations. By effective dispersal, organisms can enlarge their geographic range and adjust the dynamic, sex ratio and genetic compositions of a population. Birds are one of the groups that are studied intensively by human beings. Due to their diurnal habits, diverse life history strategies and complex movement, birds are also ideal models for the study of dispersal behaviors. Certain topics of avian dispersal including sex-biased, asymmetric dispersal caused by differences in body conditions, dispersal processes, habitat selection and long distance dispersal are discussed here. Bird-ringing or marking, radio-telemetry and genetic markers are useful tools widely applied in dispersal studies. There are three major challenges regarding theoretical study and methodology research of dispersal: (1) improvement in research methodology is needed, (2) more in-depth theoretical research is necessary, and (3) application of theoretical research into the conservation efforts for threatened birds and the management of their habitats should be carried out immediately. __________ Translated from Acta Ecologica Sinica, 2008, 28(4): 1354–1365 [译自: 生态学报]     

Social Dispersion and Genetic Variation in Two Species of Emballonurid Bats

Saccopteryx leptura and S. bilineata are closely related, sympacric species of neotropical bats that exhibit marked differences in social organization and dispersion patterns. S. leptura is monogamous and roosts in small groups of 1 to 5 (Y = 2.6) individuals that are finely dispersed. S. bilineata is harem-polygamous and roosts in larger groups of 1 to 42 (Y = 8.1) individuals that are more coarsely dispersed. Allozyme genetic studies of individuals from Trinidad, W I. demonstrate that both species carry similar, high levels of polymorphism and heterozygosity. Although significant genetic heterogeneity was observed among geographical populations of S. leptura, there is no evidence for either species that social structuring leads to inbreeding and the loss of heterozygosity, or that it promotes the development of genetic heterogeneity among social units that could accelerate the evolutionary diversification of these taxa. These results do not support a recently proposed hypothesis that social structuring in mammals has these effects. Observed intercolony genetic heterogeneity among adult males resident in multimale groups of S. bilineata is greater than intercolony heterogeneity of adult females. This is an apparent result of female dispersal and the recruitment of males into their parental groups. These results suggest that the adult males within large colonies may be kin-related.     

The spatial distribution does not affect host–parasite coevolution in <Emphasis Type="Italic">Rossomyrmex</Emphasis> ants

Host and parasite distributions are crucial to understand the coevolutionary outcomes of their relationships. This comes from the fact that the distribution of a species (fragmented vs. continuous habitats) influences its dispersal opportunities. In this work, we studied the effect of the spatial distribution on dispersal and coevolution between three species of social parasite ants of the genus Rossomyrmex (one distributed in high mountains in Spain and two distributed in extended plains in Turkey and Kazakhstan) and their ant hosts Proformica. We analysed the variation at the mitochondrial gene cytochrome c oxidase (COI) to infer female dispersal for parasites as well as the cuticular hydrocarbons (CHCs) of parasites and hosts to study their coevolutionary process, given that CHCs are involved in nestmate recognition. Our genetic results revealed a surprising scarce variation at COI for the three parasite species, suggesting selective forces that prevent from mutation fixation. Therefore, COI appeared to be a poor tool to study dispersal. Furthermore, chemical results showed population differentiation for all host–parasite systems, pointing that coevolution would take place at a local scale regardless of the spatial distribution or dispersal opportunities of the counterparts.     

Cheating does not explain selective differences at high and low relatedness in a social amoeba

Background  

Altruism can be favored by high relatedness among interactants. We tested the effect of relatedness in experimental populations of the social amoeba Dictyostelium discoideum, where altruism occurs in a starvation-induced social stage when some amoebae die to form a stalk that lifts the fertile spores above the soil facilitating dispersal. The single cells that aggregate during the social stage can be genetically diverse, which can lead to conflict over spore and stalk allocation. We mixed eight genetically distinct wild isolates and maintained twelve replicated populations at a high and a low relatedness treatment. After one and ten social generations we assessed the strain composition of the populations. We expected that some strains would be out-competed in both treatments. In addition, we expected that low relatedness might allow the persistence of social cheaters as it provides opportunity to exploit other strains.     

Dispersal of Camellia japonica seeds by Apodemus speciosus revealed by maternity analysis of plants and behavioral observation of animal vectors

Seed dispersal determines a plant’s reproductive success, range expansion, and population genetic structures. Camellia japonica, a common evergreen tree in Japan, has been the subject of recent genetic studies of population structure, but its mode of seed dispersal has been assumed, without detailed study, to be barochory. The morphological and physiological features of C. japonica seeds, which are large and nutritious, suggest zoochorous dispersal, however. We compared actual distances between mother trees and seedlings with distances attributable to gravity dispersion only, to test the zoochory hypothesis of C. japonica. The animals that transport the seeds for caching were identified experimentally. We also examined the extent to which seed dispersal is affected by the behavior of animal vectors. Seed dispersal by Apodemus speciosus was confirmed by taking photographs of animals that were consuming seeds experimentally deposited on the ground. Camellia seeds hoarded by the rodents under the litter or soil were protected from drying. On the basis of microsatellite analysis of maternal tissue from the seed coat, the mother trees of 28 seedlings were identified. Maternity analysis revealed the average seed-dispersal distance from mother trees was 5.8 m±6.0 SD, a distance greater than initial dispersal by gravity alone. These results indicate that C. japonica is a zoochorous species dispersed by A. speciosus. Fifty percent of the seed dispersal occurred from mature evergreen forests to dwarf bamboo thickets. This directional seed dispersal would contribute to range expansion of C. japonica. Home range sizes of A. speciosus were 0.85 ha at most and covered with different types of vegetation, from evergreen forests to grassland. This low specificity of their microhabitat use might enhance seed dispersal to different types of vegetation.All animal experiments complied with Japanese laws.     

Life history influences how fire affects genetic diversity in two lizard species

‘Fire mosaics’ are often maintained in landscapes to promote successional diversity in vegetation with little understanding of how this will affect ecological processes in animal populations such as dispersal, social organization and re‐establishment. To investigate these processes, we conducted a replicated, spatiotemporal landscape genetics study of two Australian woodland lizard species [Amphibolurus norrisi (Agamidae) and Ctenotus atlas (Scincidae)]. Agamids have a more complex social and territory structure than skinks, so fire might have a greater impact on their population structure and thus genetic diversity. Genetic diversity increased with time since fire in C. atlas and decreased with time since fire in A. norrisi. For C. atlas, this might reflect its increasing population size after fire, but we could not detect increased gene flow that would reduce the loss of genetic diversity through genetic drift. Using landscape resistance analyses, we found no evidence that postfire habitat succession or topography affected gene flow in either species and we were unable to distinguish between survival and immigration as modes of postfire re‐establishment. In A. norrisi, we detected female‐biased dispersal, likely reflecting its territorial social structure and polygynous mating system. The increased genetic diversity in A. norrisi in recently burnt habitat might reflect a temporary disruption of its territoriality and increased male dispersal, a hypothesis that was supported with a simulation experiment. Our results suggest that the effects of disturbance on genetic diversity will be stronger for species with territorial social organization.     

Inbreeding and soil conditions affect dispersal and components of performance of two plant species in fragmented landscapes

During habitat fragmentation, plant populations become smaller and more isolated from each other, resulting in increasing inbreeding rates within populations. Furthermore, fragmentation is often accompanied by a progressive deterioration of soil conditions. Overall, high inbreeding rates and poor soil conditions decrease plant performance and so increase the probability of extinction of fragmented plant populations. The goal of this study was to investigate the effects of inbreeding and soil acidification on seed and offspring traits of Succisa pratensis and Hypochaeris radicata, two plant species differing in mating system, lifespan and dispersal ability. For each species, plants from four populations of different sizes were hand-pollinated. The selfed and outcrossed progeny were grown at two soil pH levels. Overall, results showed that the dispersal potential of H. radicata was reduced by selfing, indicating that dispersal capacity is not independent from the genetic erosion process. Variation among seed families and its interactions with pollination treatments indicate that dispersal capacity may have a genetic basis. The performance of both species decreased sharply as soil conditions became more acidic, but inbreeding did not aggravate the process. These results suggest that S. pratensis and H. radicata populations may decline in the long term; however, family level variation suggests a potential for adaptation to new conditions.     

Frankia and Alnus rubra Canopy Roots: An Assessment of Genetic Diversity, Propagule Availability, and Effects on Soil Nitrogen

The ecological importance of microbial symbioses in terrestrial soils is widely recognized, but their role in soils that accumulate in forest canopies is almost entirely unknown. To address this gap, this study investigated the Frankia–Alnus rubra symbiosis in canopy and forest floor roots at Olympic National Park, WA, USA. Sixteen mature A. rubra trees were surveyed and Frankia genetic diversity in canopy and forest floor nodules was assessed with sequence-based nifH analyses. A seedling bioassay experiment was conducted to determine Frankia propagule availability in canopy and forest floor soils. Total soil nitrogen from both environments was also quantified. Nodules were present in the canopies of nine of the 16 trees sampled. Across the study area, Frankia canopy and forest floor assemblages were similar, with both habitats containing the same two genotypes. The composition of forest floor and canopy genotypes on the same tree was not always identical, however, suggesting that dispersal was not a strictly local phenomenon. Frankia seedling colonization was similar in canopy soils regardless of the presence of nodules as well as in forest floor soils, indicating that dispersal was not likely to be a major limiting factor. The total soil nitrogen of canopy soils was higher than that of forest floor soils, but the presence of Frankia nodules in canopy soils did not significantly alter soil nitrogen levels. Overall, this study indicates that the Frankia–A. rubra symbiosis is similar in canopy and forest floor environments. Because canopy roots are exposed to different environmental conditions within very small spatial areas and because those areas can be easily manipulated (e.g., fertilizer or watering treatments), they present microbial ecologists with a unique arena to examine root–microbe interactions.     

Genetic structure of the Black Bog Ant (<Emphasis Type="Italic">Formica picea</Emphasis> Nylander) in the United Kingdom

Social Hymenoptera have been relatively little studied in terms of conservation genetics even though their sociality and complementary sex determination potentially influence the interaction of genetics with extinction risk. Using microsatellite markers, we investigated the social and genetic structure of nests and populations of the Black Bog Ant Formica picea at four sites in the UK, where this habitat specialist has a localized and fragmented range. Nests were weakly polygynous (effective queen number, 4–27 per nest) with low worker relatedness. Isolation by distance tended to be present within sites, indicating limited dispersal, but inbreeding was rare. The four study sites fell into three main populations (two in South Wales, one in southern England). We conclude that, although UK F. picea populations are not at immediate risk from genetic factors, their limited dispersal abilities at both within- and between-site scales should inform conservation management decisions.     

Genetic Consequences of Habitat Fragmentation in Black-and-Gold Howler (<Emphasis Type="BoldItalic">Alouatta caraya</Emphasis>) Populations from Northern Argentina

Human-induced habitat fragmentation might seriously affect behavioural patterns and the survival of species whose ecological requirements strongly depend on specific environmental conditions. We compared the genetic structure and dispersal patterns of 2 populations of Alouatta caraya (Plathyrrhini, Atelidae) to understand how habitat reduction and fragmentation affect gene flow in this species. We sampled individuals from 7 groups living in continuous forest (CF, n = 46, 22 males and 24 females), and 11 groups that inhabit a fragmented forest (FF, n = 50, 24 males and 26 females). FST values based on 11 microsatellite loci showed a recent genetic differentiation among groups in the FF. In contrast, the CF showed no differentiation among groups. Further, FST values between sexes, as well as kinship relationships, also exhibited differences between habitats. In the CF, both males and females disperse, leading to nondifferentiated groups composed of adults that are not close relatives. Conversely, in the FF, some groups are differentiated, males disperse more than females, and groups are composed of closely related adult females. Our results suggest that habitat fragmentation modifies the dispersal patterns of black-and-gold howlers. These differences between habitats may reflect a reduced gene flow, providing genetic evidence that suggests that habitat fragmentation severely limits the howler’s ability to disperse. An increasing level of isolation due to uncontrolled deforestation may cause similar loss of genetic diversity on other arboreal primates, and nonprimates that depend on forest continuity to disperse, reducing their abilities to cope with environmental changes.     

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。     

Dispersal Patterns in <Emphasis Type="Italic">Tarsius spectrum</Emphasis>

One of the most widely accepted explanations for the difference in the sex bias between mammals and birds is that male-biased dispersal in mammals is due to the preponderance of polygynous mating systems exhibited by this class, whereas birds are predominantly monogamous. Spectral tarsiers (Tarsius spectrum) are unusual in that they exhibit variation in its mating system. Although the majority of spectral tarsier groups are monogamous, ca. 15% are polygynous. If mating system influences dispersal, then I predicted that the polygynous groups would exhibit male biased dispersal whereas I predicted that the dispersal patterns of the monogamous groups would be analogous to that exhibited by birds, specifically female biased. Alternatively, I hypothesized that ecological variation may influence dispersal habits in this species. Specifically, I predicted that polygynous groups would exhibit greater habitat quality than monogamous groups. The 2 hypotheses are not mutually exclusive. On the basis of 14 individuals birdbanded between 1994 and 1999, I determined that individuals of both sexes were equally likely to disperse (males, n = 5; females, n = 9). Males dispersed twice as far as females did. The mean dispersal distance for males was 660 m, and for females it was 266 m. Females (77%) were more likely to form a territory adjacent to the parental territory than were males (20%). Individuals exhibited relatively high amounts of site fidelity (86%) that were related to physical characteristics of the sleeping site. Adults that dispersed a second time (n = 4) initially resided in trees that were shorter and had a smaller diameter-at-breast height than the trees of individuals that exhibited site fidelity. The results of my study partly support the parental mating system hypothesis and also support the habitat quality hypothesis.