Kartik Shanker

Positive Relationships Between Association Strength and Phenotypic Similarity Characterize the Assembly of Mixed-Species Bird Flocks Worldwide

Competition theory predicts that local communities should consist of species that are more dissimilar than expected by chance. We find a strikingly different pattern in a multicontinent data set (55 presence-absence matrices from 24 locations) on the composition of mixed-species bird flocks, which are important subunits of local bird communities the world over. By using null models and randomization tests followed by meta-analysis, we find the association strengths of species in flocks to be strongly related to similarity in body size and foraging behavior and higher for congeneric compared with noncongeneric species pairs. Given the local spatial scales of our individual analyses, differences in the habitat preferences of species are unlikely to have caused these association patterns; the patterns observed are most likely the outcome of species interactions. Extending group-living and social-information-use theory to a heterospecific context, we discuss potential behavioral mechanisms that lead to positive interactions among similar species in flocks, as well as ways in which competition costs are reduced. Our findings highlight the need to consider positive interactions along with competition when seeking to explain community assembly.

An assessment of the olive ridley turtle (Lepidochelys olivacea) nesting population in Orissa, India

Olive ridley mass nesting events or ‘arribadas’ have been documented in Orissa, India since 1974.However, since standardised techniques have not been used to census turtles, actual population trends remain unknown.Herein, we summarise information on nesting populations in Orissa, using data from multiple sources to arrive at consensus estimates and to derive trends.We conducted a quantitative estimate of an arribada in March 1999, where nesting was estimated as � 180,000 turtles by the strip transect method. Non-linear (quadratic) fits for arribada data from 1976–1999 and a recent decrease in the size of adults suggest a potential or imminent decline, consistent with fishery-related mortality of at least 90,000 turtles since 1994.Though statistical support for the recent decline is equivocal, efforts to reduce mortality and close monitoring of the population would be prudent.The absence of reliable data on which to base conservation action highlights an urgent need to train management personnel in data collection and estimation techniques for effective monitoring of status, threats and trends. # 2003 Elsevier Ltd.All rights reserved.

The role of competition and habitat in structuring small mammal communities in a tropical montane ecosystem in southern India

Small mammals were sampled in two natural habitats (montane stunted evergreen forests and montane grassland) and four anthropogenic habitats (tea, wattle, bluegum and pine plantation) in the Upper Nilgiris in southern India. Of the species trapped, eight were in montane evergreen forests and three were in other habitats. Habitat discrimination was studied in the rodents Rattus rattus and Mus famulus and the shrew Suncus montanus in the montane forest habitat. Multivariate tests on five variables (canopy cover, midstorey density, ground cover, tree density, canopy height) showed that R. rattus uses areas of higher tree density and lower canopy cover. Suncus montanus and M. famulus use habitat with higher tree density and ground cover and lower canopy height. Multivariate tests did not discriminate habitat use between the species. Univariate tests, however, showed that M. famulus uses areas of higher tree density than R. rattus and S. montanus. Rattus rattus was the dominant species in the montane forest, comprising 60.9% of total density, while the rodent Millardia meltada was the dominant species in the grassland. Studies of spatial interaction between these two species in habitats where they coexisted showed neither overlap nor avoidance between the species. Rattus rattus, however, did use areas of lower ground cover than did M. meltada. The analysis of spatial interactions between the species, habitat discrimination and use, and the removal experiments suggest that interspecific competition may not be a strong force in structuring these small mammal communities. There are distinct patterns in the use of different habitats by some species, but microhabitat selection and segregation is weak. Other factors such as intraspecific competition may play a more important role in these communities.

From Bathymetry to Bioshields: A Review of Post-Tsunami Ecological Research in India and its Implications for Policy

More than half a decade has passed since the December 26th 2004 tsunami hit the Indian coast leaving a trail of ecological, economic and human destruction in its wake. We reviewed the coastal ecological research carried out in India in the light of the tsunami. In addition, we also briefly reviewed the ecological research in other tsunami affected countries in Asia namely Sri Lanka, Indonesia, Thailand and Maldives in order to provide a broader perspective of ecological research after tsunami. A basic search in ISI Web of Knowledge using keywords “tsunami” and “India” resulted in 127 peer reviewed journal articles, of which 39 articles were pertaining to ecological sciences. In comparison, Sri Lanka, Indonesia, Thailand and Maldives had, respectively, eight, four, 21 and two articles pertaining to ecology. In India, bioshields received the major share of scientific interest (14 out of 39) while only one study (each) was dedicated to corals, seagrasses, seaweeds and meiofauna, pointing to the paucity of research attention dedicated to these critical ecosystems. We noted that very few interdisciplinary studies looked at linkages between pure/applied sciences and the social sciences in India. In addition, there appears to be little correlation between the limited research that was done and its influence on policy in India. This review points to gap areas in ecological research in India and highlights the lessons learnt from research in other tsunami-affected countries. It also provides guidance on the links between science and policy that are required for effective coastal zone management.

Small mammal trapping in tropical montane forests of the Upper Nilgiris, southern India: an evaluation of capture-recapture models in estimating abundance.

Capture-mark-recapture was used to study small mammal populations in tropical montane forests in southern India. Eleven plots in six montane forest patches were sampled from February-October, 1994. Six species were captured, including four rodents and two shrews. PROGRAM CAPTURE was used to derive estimates of density of the most abundant species in the study area,Rattus rattus Linnaeus. The coefficient of variation of the density estimate was used as an index of precision. The coefficient of variation decreased exponentially with increasing capture probability and with an increase in trapping duration. The coefficient of variation and the capture probability were not correlated with estimates of density. The density estimate increased with trapping duration, as did trap mortality. The latter may have been due to the trend of increased mortality with recaptures of the same individual, which in turn may have been due to weight loss over consecutive captures. Estimates of density derived using four estimators were different for 2, 3, 4 and 5 days of trapping. The coefficient of variation was highest for the generalized removal estimate and lowest for Darroch’s estimate. The models and estimators could not be applied to more than one species, and for this species, only in select habitats in a few seasons. Therefore, models of density estimation developed for temperate areas may not be suitable for tropical habitats due to low densities of small mammals in these habitats.

Community structure and demography of small-mammal populations in insular montane forests in southern India

bstract Small mammals were studied in insular montane forest patches in the Upper Nilgiris in southern India from February 1994 to September 1996. Nine patches were selected at two sites, one with a single large 600-ha patch, the other with several small patches ranging in size from 0.2 to 60 ha. The population characteristics and community structure of small mammals were studied in relation to patch size and habitat structure within the patches. The two most abundant species were Rattus rattus (2–36 individuals/ha) and Suncus montanus (0–11 individuals/ha), while the abundance of seven other species recorded were very low. The population characteristics of R. rattus examined were density, biomass, proportions of juveniles, sub-adults and adults, mean weight of all animals, mean weight of adults, sex ratio and persistence. The proportion of adults, sub-adults, mean weight of animals, and mean weight of adult males were correlated with patch size. Persistence of sub-adult females in the large patch was especially high. The density and biomass of other small-mammal species were also studied. Two indices of diversity, species richness and proportion of R.␣rattus were compared as measures of community structure. Seven habitat characteristics were measured; of these, canopy cover, canopy height and tree density were correlated with the size of the patch. Density and biomass of species other than R. rattus and proportion of R. rattus were correlated with canopy height. Density and biomass of species other than R. rattus were highest in smaller patches. While the population characteristics of R. rattus may be affected by patch size, the density of rare species may be influenced by factors related to lower canopy height. Migration between patches may be an important factor in maintaining populations in these patches.

Using intra-flock association patterns to understand why birds participate in mixed-species foraging flocks in terrestrial habitats

Bird species are hypothesized to join mixed-species flocks (flocks hereon) either for direct foraging or anti-predation-related benefits. In this study, conducted in a tropical evergreen forest in the Western Ghats of India, we used intra-flock association patterns to generate a community-wide assessment of flocking benefits for different species. We assumed that individuals needed to be physically proximate to particular heterospecific individuals within flocks to obtain any direct foraging benefit (flushed prey, kleptoparasitism, copying foraging locations). Alternatively, for anti-predation benefits, physical proximity to particular heterospecifics is not required, i.e. just being in the flock vicinity can suffice. Therefore, we used choice of locations within flocks to infer whether individual species are obtaining direct foraging or anti-predation benefits. A small subset of the bird community (5/29 species), composed of all members of the sallying guild, showed non-random physical proximity to heterospecifics within flocks. All preferred associates were from non-sallying guilds, suggesting that the sallying species were likely obtaining direct foraging benefits either in the form of flushed or kleptoparasitized prey. The majority of the species (24/29) chose locations randomly with respect to heterospecifics within flocks and, thus, were likely obtaining antipredation benefits. In summary, our study indicates that direct foraging benefits are important for only a small proportion of species in flocks and that predation is likely to be the main driver of flocking for most participants. Our findings apart, our study provides methodological advances that might be useful in understanding asymmetric interactions in social groups of single and multiple species.

Glaciations, gradients, and geography: multiple drivers of diversification of bush frogs in the Western Ghats Escarpment

The historical processes underlying high diversity in tropical biodiversity hotspots like the Western Ghats of Peninsular India remain poorly understood. We sampled bush frogs on 13 massifs across the Western Ghats Escarpment and examined the relative influence of Quaternary glaciations, ecological gradients and geological processes on the spatial patterns of lineage and clade diversification. The results reveal a large in situ radiation (more than 60 lineages), exhibiting geographical structure and clade-level endemism, with two deeply divergent sister clades, North and South, highlighting the biogeographic significance of an ancient valley, the Palghat Gap. A majority of the bush frog sister lineages were isolated on adjacent massifs, and signatures of range stasis provide support for the dominance of geological processes in allopatric speciation. In situ diversification events within the montane zones (more than 1800 m) of the two highest massifs suggest a role for climate-mediated forest-grassland persistence. Independent transitions along elevational gradients among sub-clades during the Miocene point to diversification along the elevational gradient. The study highlights the evolutionary significance of massifs in the Western Ghats with the high elevations acting as centres of lineage diversification and the low- and mid-elevations of the southern regions, with deeply divergent lineages, serving as museums.

Importance of intraspecifically gregarious species in a tropical bird community

In both single- and mixed-species social groups, certain participants are known to play important roles in providing benefits. Identifying these participants is critical for understanding group dynamics, but is often difficult with large roving social groups in the wild. Here, we develop a new approach to characterize roles in social groups and apply it to mixed-species bird flocks (flocks hereafter) in an Indian tropical evergreen forest. Two types of species, namely intraspecifically gregarious and sallying species, are thought to play important roles in flocks because studies have shown they attract other flock participants. However, it is unclear why these types are attractive and whether they are essential for flock formation. We address these questions by focusing on the composition of the subset of flocks containing only two species each. In two-species flocks, it is reasonable to assume that at least one species obtains some kind of benefit. Therefore, only those species combinations that result in benefit to at least one species should occur as two-species flocks. Using data from 540 flocks overall, of which 158 were two-species flocks, we find that intraspecifically gregarious species are disproportionately represented in two-species flocks and always lead flocks when present, and that flocks containing them are joined significantly more by other species. Our results suggest that intraspecifically gregarious species are likely to be the primary benefit providers in flocks and are important for tropical flock formation. Our study also provides a new approach to understanding importance in other mixed-species and single-species social groups.

Unpacking the species conundrum: philosophy, practice and a way forward

The history of ecology and evolutionary biology is rife with attempts to define and delimit species. However, there has been confusion between concepts and criteria, which has led to discussion, debate, and conflict, eventually leading to lack of consistency in delimitation. Here, we provide a broad review of species concepts, a clarification of category versus concept, an account of the general lineage concept (GLC), and finally a way forward for species discovery and delimitation. Historically, species were considered as varieties bound together by reproduction. After over 200 years of uncertainty, Mayr attempted to bring coherence to the definition of species through the biological species concept (BSC). This has, however, received much criticism, and the last half century has spawned at least 20 other concepts. A central philosophical problem is that concepts treat species as ‘individuals’ while the criteria for categorization treats them as ‘classes’. While not getting away from this problem entirely, the GLC attempts to provide a framework where lineage divergence is influenced by a number of different factors (and correlated to different traits) which relate to the different species concepts. We also introduce an ‘inclusive’ probabilistic approach for understanding and delimiting species. Finally, we provide a Wallacean (geography related) approach to the Linnaean problem of identifying and delimiting species, particularly for cases of allopatric divergence, and map this to the GLC. Going one step further, we take a morphometric terrain approach to visualizing and understanding differences between lineages. In summary, we argue that while generalized frameworks may work well for concepts of what species are, plurality and ‘inclusive’ probabilistic approaches may work best for delimitation.