Raman Sukumar

Predicting Population Trends from Size Distributions: A Direct Test in a Tropical Tree Community

Forest ecologists often evaluate how well the species composition of saplings in the understory matches that of the canopy: absence of juveniles suggests that a tree species is suffering population decline. Here we offer a theoretical and empirical test of this assertion using data from a 50‐ha census plot in Panama. Theory indicates that higher rates of population change, λ, lead to more steeply declining size distributions (more juveniles relative to adults). But other parameters also affect the size distribution: lower growth rate of juveniles and lower survival at any size pro duce more steeply declining size distributions as well. Empirical evaluation of 216 tree populations showed that juvenile growth was the strongest predictor of size distribution, in the direction predicted by theory. Size distribution did correlate with population growth, but weakly and only in understory species, not canopy seecies. Size distribution did not correlate with the growth rate of larger individuals nor with survival. Results suggest that static in formation on the size distribution is not a good predictor of future population trends, while demographic information is. Fast‐growing species will have fewer juveniles in the understory than slow growing species, even when population growth is equal.

The Importance of Demographic Niches to Tree Diversity

Most ecological hypotheses about species coexistence hinge on species differences, but quantifying trait differences across species in diverse communities is often unfeasible. We examined the variation of demographic traits using a global tropical forest data set covering 4500 species in 10 large-scale tree inventories. With a hierarchical Bayesian approach, we quantified the distribution of mortality and growth rates of all tree species at each site. This allowed us to test the prediction that demographic differences facilitate species richness, as suggested by the theory that a tradeoff between high growth and high survival allows species to coexist. Contrary to the prediction, the most diverse forests had the least demographic variation. Although demographic differences may foster coexistence, they do not explain any of the 16-fold variation in tree species richness observed across the tropics.

Assessing Evidence for a Pervasive Alteration in Tropical Tree Communities

In Amazonian tropical forests, recent studies have reported increases in aboveground biomass and in primary productivity, as well as shifts in plant species composition favouring fast-growing species over slow-growing ones. This pervasive alteration of mature tropical forests was attributed to global environmental change, such as an increase in atmospheric CO2 concentration, nutrient deposition, temperature, drought frequency, and/or irradiance. We used standardized, repeated measurements of over 2 million trees in ten large (16–52 ha each) forest plots on three continents to evaluate the generality of these findings across tropical forests. Aboveground biomass increased at seven of our ten plots, significantly so at four plots, and showed a large decrease at a single plot. Carbon accumulation pooled across sites was significant (+0.24 MgC ha−1 y−1, 95% confidence intervals [0.07, 0.39] MgC ha−1 y−1), but lower than reported previously for Amazonia. At three sites for which we had data for multiple census intervals, we found no concerted increase in biomass gain, in conflict with the increased productivity hypothesis. Over all ten plots, the fastest-growing quartile of species gained biomass (+0.33 [0.09, 0.55] % y−1) compared with the tree community as a whole (+0.15 % y−1); however, this significant trend was due to a single plot. Biomass of slow-growing species increased significantly when calculated over all plots (+0.21 [0.02, 0.37] % y−1), and in half of our plots when calculated individually. Our results do not support the hypothesis that fast-growing species are consistently increasing in dominance in tropical tree communities. Instead, they suggest that our plots may be simultaneously recovering from past disturbances and affected by changes in resource availability. More long-term studies are necessary to clarify the contribution of global change to the functioning of tropical forests.

Ecology of the Asian Elephant in Southern India. II. Feeding Habits and Crop Raiding Patterns

The Asian elephants foraging strategy in its natural habitat and in cultivation was studied in southern India during 1981-83. Though elephants consumed at least 112 plant species in the study area, about 85% of their diet consisted of only 25 species from the order Malvales and the families Leguminosae, Palmae, Cyperaceae and Gramineae. Alteration between a predominantly browse diet during the dry season with a grass diet during the early wet season was related to the seasonally changing protein content of grasses. Crop raiding, which was sporadic during the dry season, gradually increased with more area being cultivated with the onset of rains. Raiding frequency reached a peak during October-December, with some villages being raided almost every night, when finger millet (Eleusine coracana) was cultivated by most farmers. The monthly frequency of raiding was related to the seasonal movement of elephant herds and to the size of the enclave. Of their total annual food requirement, adult bull elephants derived an estimated 9.3% and family herds 1.7% in quantity from cultivated land. Cultivated cereal and millet crops provided significantly more protein, calcium and sodium than the wild grasses. Ultimately, crop raiding can be thought of as an extension of the elephants optimal foraging strategy.

Male-female differences in foraging on crops by Asian elephants

Males and females may differ in morphology or behaviour because of contrasting factors affecting their reproductive success. In polygynous mammals with a marked sexual dimorphism, males are more likely to exhibit risky behaviour promoting reproductive success (Trivers 1985). In this study the authors present evidence that pubertal and adult male Asian elephants, Elephas maximus (above 15 years) incur greater risks than female-led family herds by foraging on cultivated crops which have more nutritive value than wild food plants.

The management of large mammals in relation to male strategies and conflict with people

Many large mammals such as elephant, rhino and tiger often come into conflict with people by destroying agricultural crops and even killing people, thus providing a deterrent to conservation efforts. The males of these polygynous species have a greater variance in reproductive success than females, leading to selection pressures favouring a ‘high risk-high gain’ strategy for promoting reproductive success. This brings them into greater conflict with people. For instance, adult male elephants are far more prone than a member of a female-led family herd to raid agricultural crops and to kill people. In polygynous species, the removal of a certain proportion of ‘surplus’ adult males is not likely to affect the fertility and growth rate of the population. Hence, this could be a management tool which would effectively reduce animal-human conflict, and at the same time maintain the viability of the population. Selective removal of males would result in a skewed sex ratio. This would reduce the ‘effective population size’ (as opposed to the total population or census number), increase the rate of genetic drift and, in small populations, lead to inbreeding depression. Plans for managing destructive mammals through the culling of males will have to ensure that the appropriate minimum size in the populations is being maintained.

Scale‐dependent relationships between tree species richness and ecosystem function in forests

1. The relationship between species richness and ecosystem function, as measured by productivity or biomass, is of long-standing theoretical and practical interest in ecology. This is especially true for forests, which represent a majority of global biomass, productivity and biodiversity.

Population differentiation within and among Asian elephant (Elephas maximus) populations in southern India

Southern India, one of the last strongholds of the endangered Asian elephant (Elephas maximus), harbours about one-fifth of the global population. We present here the first population genetic study of free-ranging Asian elephants, examining within- and among-population differentiation by analysing mitochondrial DNA (mtDNA) and nuclear microsatellite DNA differentiation across the Nilgiris-Eastern Ghats, Anamalai, and Periyar elephant reserves of southern India. Low mtDNA diversity and ‘normal’ microsatellite diversity were observed. Surprisingly, the Nilgiri population, which is the worlds single largest Asian elephant population, had only one mtDNA haplotype and lower microsatellite diversity than the two other smaller populations examined. There was almost no mtDNA or microsatellite differentiation among localities within the Nilgiris, an area of about 15 000 km2. This suggests extensive gene flow in the past, which is compatible with the home ranges of several hundred square kilometres of elephants in southern India. Conversely, the Nilgiri population is genetically distinct at both mitochondrial and microsatellite markers from the two more southerly populations, Anamalai and Periyar, which in turn are not genetically differentiated from each other. The more southerly populations are separated from the Nilgiris by only a 40-km-wide stretch across a gap in the Western Ghats mountain range. These results variably indicate the importance of population bottlenecks, social organization, and biogeographic barriers in shaping the distribution of genetic variation among Asian elephant populations in southern India.

The line transect method for estimating densities of large mammals in a tropical deciduous forest: An evaluation of models and field experiments

We have evaluated techniques of estimating animal density through direct counts using line transects during 1988–92 in the tropical deciduous forests of Mudumalui Sanctuary in southern India for four species of large herbivorous mammals, namely, chital (Axis axis). sambar (Cervus unicolor). Asian elephant (Elephas maximus) and gaur (Bos gaurus)Density estimates derived from the Fourier Series and the Half-Normal models consistently had the lowest coefficient of variation. These two models also generated similar mean density estimates. For the Fourier Series estimator, appropriate cut-off widths for analyzing line transect data for the four species are suggested. Grouping data into various distance classes did not produce any appreciable differences in estimates of mean density or their variances, although model fit is generally better when data arc placed in fewer groups. The sampling effort needed to achieve a desired precision (coefficient of variation) in the density estimate is derived. A sampling effort of 800 km of transects returned a 10% coefficient of variation on estimate for ehital; for the other species a higher effort was needed to achieve this level of precision. There was no statistically significant relationship between detectability of a group and the size of the group for any species. Density estimates along roads were generally significantly different from those in the interior of the forest, indicating that road-side counts many not be appropriate for most species.

Social organization of the Asian elephant ( Elephas maximus ) in southern India inferred from microsatellite DNA

Social organization of the Asian elephant (Elephas maximus) is not well understood in the absence of long-term studies of identified individuals. Adult Asian elephant females and their young offspring of both sexes form matriarchal groups, with pubertal males dispersing from natal groups, but whether these social groups represent families and whether males show locational or social dispersal were unknown. Using nuclear microsatellite loci amplified from dung-extracted DNA of free-ranging elephants in a large southern Indian population, we demonstrate that female-led herds comprise closely related individuals that are indeed families, and that males exhibit non-random locational dispersal.