Warren Y. Brockelman

An Analysis of Density Effects and Predation in Bufo Americanus Tadpoles

The effects of density on growth and survivorship of Bufo americanus tadpoles and the importance of some predators were investigated experimentally in two small ponds in southeastern Michigan. Tadpoles were raised through metamorphosis at different densities and food levels in screened inclosures placed in the ponds. Most inclosures were open to the natural substratum but others were completely screened to exclude larger predators. In one pond experiment, time of metamorphosis, individual growth variability and mortality were directly related to initial tadpole density, and transformation size was inversely related. The addition of food (natural periphyton on plant litter) produced a density—dependent increase in growth rate. The density—related mortality was correlated with variations in growth rate among pens. Growth rate was correlated with submerged plant cover in the pens. Neither survivorship nor growth was correlated with predator densities. The other pond experiment was characterized by a lack of ...

Dispersal, pair formation and social structure in gibbons (Hylobates lar)

Abstract We report observations on reproduction, natal dispersal, pair formation, and group structure based on longitudinal observations of several white-handed gibbon (Hylobates lar) groups spanning 18 years. Our observations are at odds with the traditional view that gibbons live in nuclear family groups consisting of a pair of adults and their offspring, and that parents exclude young from the family territory when they reach adult size. In the relatively dense Khao Yai study population, dispersing young usually obtain mates by replacing adults in existing territories, which creates non-nuclear families. Six subadults, five males and one female, matured and dispersed at an average age of 10 years, or about 2 years after reaching adult size. Average natal dispersal distance was 710 m, or between one and two territories away. At least two dispersing males replaced adults in neighboring groups. In one case, forcible displacement of the resident male resulted in a group which included a young juvenile presumably fathered by the previous male, two younger juveniles (probably brothers) from the new males original group, and (later) offspring of the new pair. Social relations within this heterogeneous group remained harmonious: the adults groomed all the young and play occurred between all preadult members. In only two out of a total of seven cases of dispersal seen did two subadults pair and disperse into new territorial space. Nonreproducing subadults which delay dispersal may be tolerated by the adults provided that they contribute benefits to the adults or their offspring. Possible benefits include behaviors such as grooming, social play with juveniles, and support of the adult male in defending the territory. Delayed dispersal is probably advantageous in a saturated environment where there is no room for floaters, but subadults may also gain indirect fitness benefits by aiding siblings and other relatives.

Bushmeat poaching reduces the seed dispersal and population growth rate of a mammal-dispersed tree

Myriad tropical vertebrates are threatened by overharvest. Whether this harvest has indirect effects on nonhunted organisms that interact with the game species is a critical question. Many tropical birds and mammals disperse seeds. Their overhunting in forests can cause zoochorous trees to suffer from reduced seed dispersal. Yet how these reductions in seed dispersal influence tree abundance and population dynamics remains unclear. Reproductive parameters in long-lived organisms often have very low elasticities; indeed the demographic importance of seed dispersal is an open question. We asked how variation in hunting pressure across four national parks with seasonal forest in northern Thailand influenced the relative abundance of gibbons, muntjac deer, and sambar deer, the sole dispersers of seeds of the canopy tree Choerospondias axillaris. We quantified how variation in disperser numbers affected C. axillaris seed dispersal and seedling abundance across the four parks. We then used these data in a structured population model based on vital rates measured in Khao Yai National Park (where poaching pressure is minimal) to explore how variation in illegal hunting pressure might influence C. axillaris population growth and persistence. Densities of the mammals varied strongly across the parks, from relatively high in Khao Yai to essentially zero in Doi Suthep-Pui. Levels of C. axillaris seed dispersal and seedling abundance positively tracked mammal density. If hunting in Khao Yai were to increase to the levels seen in the other parks, C. axillaris population growth rate would decline, but only slightly. Extinction of C. axillaris is a real possibility, but may take many decades. Recent and ongoing extirpations of vertebrates in many tropical forests could be creating an extinction debt for zoochorous trees whose vulnerability is belied by their current abundance.

Field studies on the transmission of the human liver fluke, Opisthorchis viverrini, in northeast Thailand: population changes of the snail intermediate host.

Abstract Field studies on the transmission of the human liver fluke, Opisthorchis viverrini , in northeast Thailand: population changes of the snail intermediate host. International Journal for Parasitology 16 : 545–552. A natural population of Bithynia siamensis goniomphalos , snail vector of the human liver fluke Opisthorchis viverrini was sampled for 3 years in a shallow reservoir in Khon Kaen Province, northeast Thailand. During the study period, the habitat experienced a 17-month drought followed by a surging flood. Snails suffered over 90% mortality during drying of the reservoir but died at an average rate of only 5% per month during aestivation. When water returned they migrated to new edge habitats, apparently on floating debris, and reproduced. Peaks in reproduction occurred following spring rains and after the fall monsoon flooding subsided. Normally, two generations are produced per year in seasonal habitats. The prevalence of O. viverrini infection averaged 0.11 % in adult snails over 8 mm in length; smaller snails were not infected. Control of opisthorchiasis through snail control does not appear practical because of the widespread distribution of the snails, their ability to survive in very unstable habitats, and for other reasons concerned with the existence of the fish host between snail and human hosts.

Rate of re-infection by Opisthorchis viverrini in an endemic Northeast Thai community after chemotherapy

Patterns of re-infection by the human liver fluke, Opisthorchis viverrini, were studied over a period of 1 year after chemotherapy in a community in Northeast Thailand. Praziquantel (40 mg per kg body weight) was administered to 808 subjects; 88.4% of those examined after 2 weeks had negative stools. Within 1 year of treatment, however, 87.7% of the cured subjects who were re-examined had become re-infected, and 51.5% had infections of at least moderate intensity (1 epmg of faeces), in comparison with 72% of the same group before treatment. Those with high pre-treatment intensities of infection tended to have heavier intensities of re-infection, indicating that some people are predisposed to heavy infections. The rate of re-infection of those with heavy pre-treatment intensity of infection was about twice that of those who were negative or had only light infection before treatment. The rate of re-infection was markedly higher than concurrently and previously measured natural incidences of infection. These findings suggest that chemotherapy would have to be applied several times a year in order to control opisthorchiasis, and that it might be most cost-effective to preferentially treat heavily infected individuals.

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.

Functional differences within a guild of tropical mammalian frugivores

Many plants interact with groups of mutualist pollinators and seed dispersers. A key issue for both basic ecology and conservation is whether the different species within these guilds of mutualist animals are functionally equivalent. Comparing the relative effects of sympatric mutualists is important for understanding the evolution of multispecies mutualisms and for predicting mutualism stability in the face of anthropogenic change. However, empirical comparisons of the population-level impacts of mutualist animals on their host plant are rare, particularly for seed dispersal mutualisms in species-rich ecosystems. We compared the influence of three seed-dispersing tropical mammals, lar gibbons (Hylobates lar), sambar deer (Rusa unicolor), and red muntjac deer (Muntiacus muntjak), on the demography of a shared host tree in Thailand, Choerospondias axillaris (Anacardiaceae). Sambar and muntjac dispersed far more C. axillaris seeds than did gibbons. While sambar deposited many seeds under female tree canopies, muntjac were the only disperser to move seeds to open microhabitats, where C. axillaris seed germination, seedling survival, and initial growth are enhanced. Using stage-based population models, we assessed how disperser-specific seed dispersal, variation in the frequency of canopy gap formation, and their interaction influenced the potential population growth of C. axillaris. Large differences in dispersal quantity and small differences in dispersal quality among sambar and gibbons resulted in similar and negligible impacts on the trees population dynamics. Muntjac, by taking some of the seeds to open microhabitats, are projected to have a greater positive impact on C. axillaris demography than either sambar or gibbons. Model comparisons of population-level species impacts may allow us to predict which ecological interactions are at risk from loss of critical species.

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.

A Field Test of Distance Sampling Methods for a Tropical Forest Bird Community

ABSTRACT. Distance sampling is usually preferred over uncorrected point counts for surveys of forest birds, but rarely has its accuracy been assessed against known numbers, particularly in tropical forests. We compared density estimates of eight species of breeding bird—Black-naped Monarch (Hypothymis azurea), Hill Blue Flycatcher (Cyornis banyumas), White-rumped Shama (Copsychus malabaricus), Puff-throated Bulbul (Alophoixus pallidus), Abbotts Babbler (Malacocincla abbotti), Puff-throated Babbler (Pellorneum ruficeps), White-browed Scimitar Babbler (Pomatorhinus schisticeps), and White-bellied Yuhina (Yuhina zantholeuca)—obtained through color-banding, nest finding, and territory mapping with those derived from distance methods in evergreen forest in northeastern Thailand. We also assessed the availability of birds to be detected using a closed-capture model and incorporated this with point-transect distance sampling. Abundance estimates from territory mapping and distance sampling were highly correlated, but biased for two species using line transects and five species using point transects. Six of the seven biased estimates were biased low. Probabilities of detection were not significantly different between lines and points, and there was no significant difference in the overall accuracy between methods. Accounting for observer differences improved density estimates but reduced precision. The variance in accuracy was mostly related to the behavior of the different species. Adjusting for availability did not improve the overall accuracy of the estimates, because of the low singing rates of tropical birds. Nonetheless, distance sampling provided relatively robust estimates despite the near total dependence on aural cues. Violations of distance-sampling assumptions may be frequent in heavily forested habitats, where both availability for detection and probability of detection on the transect line (or point) are likely to be <1.

Plant diversity increases with the strength of negative density dependence at the global scale

Maintaining tree diversity Negative interaction among plant species is known as conspecific negative density dependence (CNDD). This ecological pattern is thought to maintain higher species diversity in the tropics. LaManna et al. tested this hypothesis by comparing how tree species diversity changes with the intensity of local biotic interactions in tropical and temperate latitudes (see the Perspective by Comita). Stronger local specialized biotic interactions seem to prevent erosion of biodiversity in tropical forests, not only by limiting populations of common species, but also by strongly stabilizing populations of rare species, which tend to show higher CNDD in the tropics. Science, this issue p. 1389; see also p. 1328 A global analysis of ~3000 species and ~2.4 million trees elucidates variations in tree species diversity between tropical and temperate latitudes. Theory predicts that higher biodiversity in the tropics is maintained by specialized interactions among plants and their natural enemies that result in conspecific negative density dependence (CNDD). By using more than 3000 species and nearly 2.4 million trees across 24 forest plots worldwide, we show that global patterns in tree species diversity reflect not only stronger CNDD at tropical versus temperate latitudes but also a latitudinal shift in the relationship between CNDD and species abundance. CNDD was stronger for rare species at tropical versus temperate latitudes, potentially causing the persistence of greater numbers of rare species in the tropics. Our study reveals fundamental differences in the nature of local-scale biotic interactions that contribute to the maintenance of species diversity across temperate and tropical communities.