The ecology of white-handed and pileated gibbons in a zone of overlap and hybridization in Thailand.

OBJECTIVES: The study of related species in contact zones can elucidate what factors mediate species coexistence and geographical distributions. We investigated niche overlap and group interactions of two gibbon species and their hybrids co-occurring in a zone of overlap and hybridization. METHODS: The location, composition and behavior of white-handed, pileated, and mixed-species gibbon groups were studied by following them during 31 consecutive months in a relatively large part of the contact zone. RESULTS: Twenty groups of white-handed gibbon were mapped followed by nine groups of pileated gibbons and five mixed-species groups. White-handed, pileated and mixed-species groups had similar sizes and composition, ate a high proportion of fruits, shared a large number of species in their diets, and presented similar habitat preferences. Group home range sizes did not differ between species and overlapped little with neighboring groups irrespective of species, and intraspecific and interspecific encounter rates were similar. DISCUSSION: Ecological similarities support that competition between the gibbon species exists and takes the form of interspecific territoriality. However, we could not find any clear mechanism of niche partitioning favoring coexistence between species. Our findings suggest that the contact zone is unstable and is maintained by dispersal inward from groups of the parental species. The relatively low numbers of mixed-species groups and hybrids found suggests a high degree of premating reproductive isolation, perhaps mediated by interspecific miscommunication. The existence of hybrids and backcrosses potentially undetectable from phenotypic characters alone raises the possibility of more widespread introgression than has been evident. Hence, while interspecific territoriality should reduce the rate of gene transfer, it would not necessarily present a barrier to introgression into contiguous populations of the opposite species.

Ranging and site fidelity in northern pigtailed macaques (Macaca leonina) over different temporal scales.

Space-use patterns are crucial to understanding the ecology, evolution, and conservation of primates, but detailed ranging data are scarce for many species, especially those in Southeast Asia. Researchers studying site fidelity to either home ranges or core areas have focused mainly on territorial species, whereas less information is available for non-territorial species. We analyzed the ranging patterns and site fidelity of one wild troop of northern pigtailed macaques over 16 months at different temporal scales. We used characteristic hull polygons in combination with spatial statistics to estimate home ranges and core areas. The total home range and core areas were 449 ha and 190 ha, respectively. Average daily path length was 2,246 m. The macaques showed a high defendabili--ty index according to the expected ranging of a non-territorial species in which movement does not theoretically permit the defense of a large territory. Overall, the study troop ranged more extensively than conspecific groups and closely related species studied elsewhere. These differences may reflect variable troop size, degree of terrestriality and habitat characteristics, but could also reflect methodological differences. The location, size and shape of home ranges and core areas, and extent of daily path lengths changed on a monthly basis resulting in low site fidelity between months. The macaques also showed clear shifts in the location of daily home ranges with low site fidelity scores between consecutive days. Daily home range and daily path length were related to seasonality, with greater values during the fruit-abundant period. Low site fidelity associated with lack of territoriality is consistent with macaques structuring their movement based on available food sources. However, ranging patterns and site fidelity can also be explained by macaques feeding on the move, a foraging strategy that hinders frequent and long visits to the same location.

Quality and overlap of individual core areas are related to group tenure in female spider monkeys.

In species with a high degree of fission-fusion dynamics group members may differ in the use of the group home range to reduce food competition. Such differential use may result in distinct individual core areas. We studied core area quality and overlap among 21 female spider monkeys belonging to the same group over a period of 4 years. Core areas ranged between 62 and 161 ha with a mean overlap of 56% between any given two females. Only a small portion (mean = 3 ha) of each individual core area was used exclusively. No single part of the home range was used as core area by all females, and only an area of less than 1 ha was used as part of the core area by 20 of the 21 females. The time a female spent in the group (i.e., group tenure) was associated with characteristics of the core areas: the longer the group tenure, the better the quality of her core area. In addition, the longer the time two females spent together in the same group, the larger the overlap between their individual core areas. As this result was obtained while controlling for the time two females spent together in the same subgroup, females may reduce direct competition by using the same resource at different times. In sum, spider monkey females' group tenure plays a central role in the quality and overlapping patterns of their individual core areas.

Genetic analysis of hybridization between white-handed (Hylobates lar) and pileated (Hylobates pileatus) gibbons in a contact zone in Khao Yai National Park, Thailand.

Natural hybridization has played various roles in the evolutionary history of primates. Its consequences range from genetic introgression between taxa, formation of hybrid zones, and formation of new lineages. Hylobates lar, the white-handed gibbon, and Hylobates pileatus, the pileated gibbon, are largely allopatric species in Southeast Asia with a narrow contact zone in Khao Yai National Park, Thailand, which contains both parental types and hybrids. Hybrid individuals in the zone are recognizable by their intermediate pelage and vocal patterns, but have not been analyzed genetically. We analyzed mitochondrial and microsatellite DNA of 52 individuals to estimate the relative genetic contributions of the parental species to each individual, and the amount of introgression into the parental species. We obtained fecal samples from 33 H. lar, 15 H. pileatus and four phenotypically intermediate individuals in the contact zone. Both mitochondrial and microsatellite markers confirmed distinct differences between these taxa. Both H. lar and H. pileatus contributed to the maternal lineages of the hybrids based on mitochondrial analysis; hybrids were viable and present in socially normal reproductive pairs. The microsatellite analysis identified ten admixed individuals, four F1 hybrids, which corresponded to phenotypic hybrids, and six H. lar-like backcrosses. All 15 H. pileatus samples were identified as originating from genetically H. pileatus individuals with no H. lar admixture; hence, backcrossing is biased toward H. lar. A relatively low number of phenotypic hybrids and backcrossed individuals along with a high number of parental types indicates a bimodal hybrid zone, which suggests relatively strong bias in mate selection between the species.

Gibbon travel paths are goal oriented.

Remembering locations of food resources is critical for animal survival. Gibbons are territorial primates which regularly travel through small and stable home ranges in search of preferred, limited and patchily distributed resources (primarily ripe fruit). They are predicted to profit from an ability to memorize the spatial characteristics of their home range and may increase their foraging efficiency by using a 'cognitive map' either with Euclidean or with topological properties. We collected ranging and feeding data from 11 gibbon groups (Hylobates lar) to test their navigation skills and to better understand gibbons' 'spatial intelligence'. We calculated the locations at which significant travel direction changes occurred using the change-point direction test and found that these locations primarily coincided with preferred fruit sources. Within the limits of biologically realistic visibility distances observed, gibbon travel paths were more efficient in detecting known preferred food sources than a heuristic travel model based on straight travel paths in random directions. Because consecutive travel change-points were far from the gibbons' sight, planned movement between preferred food sources was the most parsimonious explanation for the observed travel patterns. Gibbon travel appears to connect preferred food sources as expected under the assumption of a good mental representation of the most relevant sources in a large-scale space.

Interspecific interaction and predator avoidance behavior in response to tayra (Eira barbara) by mantled howler monkeys (Alouatta palliata).

This paper reports the response of one howler monkey group (Alouatta palliata) to a group of potential predators, the tayra (Eira barbara). The apparently successful predator avoidance behavior of the monkeys was recorded in detail. We observed a group of four adult tayras moving around the Alouatta group displaying a species-typical aggressive behavioral pattern. The two adult females of the howler group successfully chased the tayras away by repeatedly moving closer to the mustelids and even following them until the predators moved off.

Variability in core areas of spider monkeys (Ateles geoffroyi) in a tropical dry forest in Costa Rica.

Core areas are highly used parts of the home range on which the survival of solitary or group-living animals depends. We investigated the home range and core area size and area fidelity of a spider monkey community in a tropical dry forest over a 4-year period. Home ranges overlapped extensively across years, subgroup sizes, and seasons. In contrast, spider monkeys used core areas that varied in size and location across the study years, subgroup sizes, and seasons. These shifts in core areas suggest that the understanding of core areas, and thus the spatial requirements, of a species in a particular habitat may be limited if based on short-term studies. In this respect, our findings emphasize the importance of long-term studies of the spatial ecology of any species in a particular habitat. Our study also shows that the yearly home range basically includes all the core areas from different years, seasons, and subgroup sizes (i.e., the super-core area). This is conceptually important for territorial species, such as spider monkeys, which defend a stable home range as it contains not only the current, but also the future core areas.

Demography, life history and migrations in a Mexican mantled howler group in a rainforest fragment.

This paper represents the results of a long-term study (1996-2003) on the demographic changes over time of a Mexican mantled howler (Alouatta palliata mexicana) group in a rainforest fragment (40 ha) in Los Tuxtlas, Mexico, with a follow-up census 3 years later (2006). In addition to demographic and life history parameters, we describe six dispersal events. Our results suggest that this group has been expanding during the study period, growing from six to 12 individuals, with an annual average intrinsic growth rate of 0.07, an infant survivorship of 67%, and an average immature to female ratio of 0.90. This increase in size is probably related to the high food availability in their home range. However, fragment isolation may be negatively affecting the dispersal patterns typical of the species, which could result in a loss of genetic variability over time.

Foraging habits of Alouatta palliata mexicana in three forest fragments.

The activity patterns and diet of howler monkeys (Alouatta palliata mexicana) were studied in 3 forest fragments in south-eastern Mexico: Playa Escondida (PLA), Agaltepec Island (AGA) and Arroyo Liza (LIZ). Intersite differences offered the opportunity to investigate the foraging adaptations of howler monkeys in response to population and habitat size. In the largest fragment (PLA), the howlers' diet was based on high-quality items (fruit and young leaves). In AGA, where the density of howlers was the highest, their diet was mostly folivorous with a marked exploitation of uncommon food items such as vines, lianas, shrubs and herbs. The dietary differences in AGA were accompanied by more time spent travelling and less time spent resting. Although LIZ was the smallest fragment and had a high howler density, the small group size and the use of energy-minimizing strategies (less time spent travelling and more time spent resting) probably allowed howlers to maintain a frugivorous diet.

Biogeographical and floristic predictors of the presence and abundance of mantled howlers (Alouatta palliata mexicana) in rainforest fragments at Los Tuxtlas, Mexico.

This research focuses on identifying the principal habitat characteristics that influence the presence and abundance of mantled howlers in forest fragments. We provide information on the demography of several fragmented Alouatta palliata mexicana subpopulations at Los Tuxtlas, Mexico, and relate this to the biogeographical and floristic characteristics of the forest fragments inhabited. The most important habitat characteristics related to the presence and abundance of howlers in the fragments were fragment size and floristic diversity. On the other hand, some evidence suggests that given the conditions under which howlers in our study area live (i.e., small and degraded fragments with high densities), secondary vegetation may be beneficial for the survival of the howlers. Finally, we discuss the possibility that the very low immature-to-female ratio (IFR) in the groups, and the lack of juveniles found in many of the study groups may be due to high mortality rates in immatures. A reduction in food availability because of the high population densities of these groups may be responsible for this process.