Multispecies coexistence of trees in tropical forests: spatial signals of topographic niche differentiation increase with environmental heterogeneity

Neutral and niche theories give contrasting explanations for the maintenance of tropical tree species diversity. Both have some empirical support, but methods to disentangle their effects have not yet been developed. We applied a statistical measure of spatial structure to data from 14 large tropical forest plots to test a prediction of niche theory that is incompatible with neutral theory: that species in heterogeneous environments should separate out in space according to their niche preferences. We chose plots across a range of topographic heterogeneity, and tested whether pairwise spatial associations among species were more variable in more heterogeneous sites. We found strong support for this prediction, based on a strong positive relationship between variance in the spatial structure of species pairs and topographic heterogeneity across sites. We interpret this pattern as evidence of pervasive niche differentiation, which increases in importance with increasing environmental heterogeneity.     

Estimation of density of gibbon groups by use of loud songs

The density of gibbon populations may be estimated by listening for the loud duetted songs of monogamous territorial groups. This method requires a correction factor which must be estimated from the frequency of singing of an adequate number of known study groups. The correction factor and its error were estimated for pileated gibbons (Hylobates pileatus) in Khao Soi Dao Wildlife Sanctuary in southeastern Thailand. Among 30 groups studied, 47% sang per day, on average, but the variation between days and the variation in singing frequency between groups were large. Weather conditions, especially windiness, explained some variation in singing. During an area-wide survey of groups in the sanctuary, unexplained variation in singing from day to day accounted for approximately half of the sample error of group density estimated from 1-day listening samples. Error due to day-to-day variability can be reduced by listening for more than one day at each site. Correction factors based on the cumulative proportion of groups heard during longer (2–5-day) sample periods of listening were closer to 1.0, therefore leaving less room for error and bias of the correction factor. © 1993 Wiley-Liss, Inc.     

Bed site selection of red muntjac (<Emphasis Type="Italic">Muntiacus muntjak</Emphasis>) and sambar (<Emphasis Type="Italic">Rusa unicolor</Emphasis>) in a tropical seasonal forest

The selection of bedding sites is important for the ecology of ruminants, but has mainly been described for temperate species. Here we assessed the bed site selection of two Southeast Asian tropical deer, red muntjac and sambar, in Khao Yai National Park, Thailand. We surveyed transects weekly for 10 weeks each in 2003 and 2004 to locate bed sites, and compared the slope, aspect, and forest canopy cover of bed site locations between the two species and with available habitat. As with most temperate deer, muntjac and sambar both avoided sites with low levels of cover for their bed site locations; this could be for concealment or thermoregulation. Sambar also selected flatter sites than would be expected by the availability of topographic slopes; this could be to reduce the energy associated with getting to and from bed sites, or to increase long-range visibility from sites. Muntjac and sambar differed in their choice of aspects for bed sites; muntjac disproportionately chose west-facing areas, while sambar chose east-facing locations. This could represent a strategy by which one species avoids the other, or else differential resource requirements between the two species.     

Pileated Gibbon Density in Relation to Habitat Characteristics and Post‐logging Forest Recovery

ABSTRACT Although it is known that forest loss and degradation negatively impact most forest‐dwelling primates, such relationships are difficult to quantify because many primates are difficult to survey over large areas. Furthermore, recovery times are also difficult to assess due to a lack of long‐term data. Here, we determined how forest characteristics and habitat disturbance correlate with the abundance of pileated gibbons, Hylobates pileatus. We studied a population in Khao Ang Rue Nai Wildlife Sanctuary in southeastern Thailand, assessed its density using an auditory method combined with distance sampling at 24 randomly placed sample sites. In addition, we determined how simple forest structural characteristics and habitat disturbance correlate with the gibbon abundance. Average gibbon density per site was 1.02 ± 0.16 (SE) groups/km² (range 0–2.74). Bivariate analyses indicated that densities depended on food tree biomass, level of disturbance, evergreen forest cover, time since protection, and distance to the sanctuary boundary. Multiple regression analysis suggested evergreen forest cover and distance to boundary were the most influential factors. Because evergreen forest cover, time since protection, and habitat disturbance are correlated, these results suggest a direct dependence of gibbon densities on mature, undisturbed evergreen forest. While gibbons can persist in disturbed areas if the forest is protected, it appears that recovery to previous densities may take decades. We suggest that this is due to the slow pace of forest regeneration and/or poor recovery potential of gibbons.     

Infant loss during and after male replacement in gibbons

According to the sexual selection hypothesis, infanticide during resident male replacement is an adaptive strategy that has evolved because the killing of unweaned offspring sired by previous males shortens the inter‐birth intervals of the mothers whose infants are targeted and thereby increases the reproductive fitness of the perpetrator. To test this hypothesis, we describe previously unreported cases of primary male replacement for two gibbon species (Hylobates lar and Nomascus nasutus), and review all other reported cases of primary male replacement in gibbons. Overall, infants were present in nearly half of all cases (16/33, 48%) and of the 18 infants present during replacement, 50% (N = 9) disappeared within 2 months of the event. In four of the five cases where there was sufficient demographic information to identify the likely sire of the subsequent offspring of females that lost infants, the new male was believed to be the sire. Infants were also less likely to die or disappear if the new male and original resident male were possible kin. However, there was no significant difference in the age of infants between those that died or disappeared following replacement and those that survived to weaning (p = .630). Our review of takeover‐related infant loss in gibbons confirms that periods of male instability are risky for unweaned infants and that replacing males benefit from infant loss. Nevertheless, variability in the context of infant loss and difficulties related to data collection in the field make it difficult to test competing hypotheses concerning the mechanisms and functions of infanticide in the small apes.     

Social correlates of reproductive success in the gibbon colony on Ko Klet Kaeo, Thailand

The free-ranging colony of gibbons established on an island in the Gulf of Thailand by the SEATO Medical Research Laboratory in 1966–1967 was terminated in 1970. Out of the 20 gibbons originally introduced, four adults of each sex remained until near the end of the project. The four females each gave birth to one infant. In all cases the young were conceived five or more months after pairbonds and ranges had stabilized into a more natural-like pattern, with fairly regular morning inter-group vocal sessions and territorial boundaries between groups containing the most feral individuals. The maximum effective breeding density on the 60-acre island was found to be about four or five pairs. The chances of reproductive success appear to be maximized in a free-ranging colony containing several pairs by maintaining a one-to-one sex ratio and allowing enough space so that natural patterns of social behavior may be expressed. The amount of area required per group may vary with the individual gibbons and the environment, but we found it to be about ten acres.     

Different responses of soil respiration to environmental factors across forest stages in a Southeast Asian forest

Soil respiration (SR) in forests contributes significant carbon dioxide emissions from terrestrial ecosystems and is highly sensitive to environmental changes, including soil temperature, soil moisture, microbial community, surface litter, and vegetation type. Indeed, a small change in SR may have large impacts on the global carbon balance, further influencing feedbacks to climate change. Thus, detailed characterization of SR responses to changes in environmental conditions is needed to accurately estimate carbon dioxide emissions from forest ecosystems. However, data for such analyses are still limited, especially in tropical forests of Southeast Asia where various stages of forest succession exist due to previous land‐use changes. In this study, we measured SR and some environmental factors including soil temperature (ST), soil moisture (SM), and organic matter content (OM) in three successional tropical forests in both wet and dry periods. We also analyzed the relationships between SR and these environmental variables. Results showed that SR was higher in the wet period and in older forests. Although no response of SR to ST was found in younger forest stages, SR of the old‐growth forest significantly responded to ST, plausibly due to the nonuniform forest structure, including gaps, that resulted in a wide range of ST. Across forest stages, SM was the limiting factor for SR in the wet period, whereas SR significantly varied with OM in the dry period. Overall, our results indicated that the responses of SR to environmental factors varied temporally and across forest succession. Nevertheless, these findings are still preliminary and call for detailed investigations on SR and its variations with environmental factors in Southeast Asian tropical forests where patches of successional stages dominate.     

Behavioral and Social Structure Effects on Seed Dispersal Curves of a Forest‐Interior Bulbul (Pycnonotidae) in a Tropical Evergreen Forest

Movement patterns of animals can vary dramatically as a function of their reproductive cycle or social structure; however, little is known about how changes in the social structure of dispersers affect patterns of seed dispersal. We examined the movement patterns of the forest‐dwelling and cooperatively breeding Puff‐throated Bulbul (Alophoixus pallidus) in relation to different stages of their reproductive cycle, time of day, and group size, to determine potential impacts on the shape and scale of dispersal curves generated using a combination of gut passage time and displacement distance data. There were significant differences in dispersal distances depending on group size, season (breeding, non‐breeding), incubation (vs. other times of the year), and time of the day. The estimated median seed dispersal distance was 28 m. The median dispersal distances produced by birds in larger groups were longer than those of smaller groups (29 m vs. 25 m). During the breeding season, median dispersal distances were longer than during the non ‐ breeding season (31 m vs. 25 m), but the median dispersal distances were significantly shorter during incubation than during outside incubation (24 m vs. 28 m). The median dispersal distance produced in the early morning (30 m) was also longer than that of other times of the day (23 m late morning, 28 m early afternoon, and 26 m late afternoon). This study suggests that various aspects of an animal's behavior are likely to have significant effects on seed shadows and that this may vary significantly even among individuals of the same species.     

Interspecific territoriality in gibbons (Hylobates lar and H. pileatus) and its effects on the dynamics of interspecies contact zones

We investigated the ecology and interspecific interactions of the two gibbon species (Hylobates lar and H. pileatus) that overlap in distribution within a narrow zone of contact in the headwaters of the Takhong River at Khao Yai National Park in central Thailand. The zone is about 10-km wide, with phenotypic hybrids comprising 6.5% of the adult population. We compared species with respect to diet, territory size, intra- and interspecific group encounters, and territory quality. The two gibbon species exploited the same types of resources within their territories despite variation in the relative abundance of food-plant species between territories. The gibbons were interspecifically territorial, and males of both species displayed aggressive behaviors at both intraspecific and interspecific territorial boundaries. There was no marked difference in the amount of overlap between territories of conspecific and heterospecific pairs of groups. Although the habitat was not homogeneous, territory quality did not vary significantly between species. The species have not diverged in habitat preference or in diet. Neither species dominated in interspecific encounters, and both were reproducing well in the contact zone. We analyzed the potential consequences of several types of interspecific interactions on individual dispersal options and on the structure of the contact zone. Interference competition through interspecific territoriality affects the dispersal of individuals into the range of the other species. In general, territorial competition coupled with limited hybridization leads to predictions of a narrow contact zone or parapatry between species; thus, behavioral and ecological interactions between species need to be considered as potential factors in explaining range borders of primate species.