Ecology of the Asian Elephant in Southern India. I. Movement and Habitat Utilization Patterns

The movement and habitat utilization patterns were studied in an Asian elephant population during 1981-83 within a 1130 km2 area in southern India (110 30' N to 120 0' N and 760 50' E to 770 15' E). The study area encompasses a diversity of vegetation types from dry thorn forest (250-400 m) through deciduous forest (400-1400 m) to stunted evergreen shola forest and grassland (1400-1800 m). Home range sizes of some identified elephants were between 105 and 320 km2. Based on the dry season distribution, five different elephant clans, each consisting of between 50 and 200 individuals and having overlapping home ranges, could be defined within the study area. Seaso- nal habitat preferences were related to the availability of water and the palatability of food plants. During the dry months (January-April) elephants congregated at high densities of up to five individuals kM-2 in river valleys where browse plants had a much higher protein content than the coarse tall grasses on hill slopes. With the onset of rains of the first wet season (May- August) they dispersed over a wider area at lower densities, largely into the tall grass forests, to feed on the fresh grasses, which then had a high protein value. During the second wet season (September-December), when the tall grasses became fibrous, they moved into lower elevation short grass open forests. The normal movement pattern could be upset during years of adverse environmental con- ditions. However, the movement pattern of elephants in this region has not basically changed for over a century, as inferred from descriptions recorded during the nineteenth century.

Growth in the Asian elephant

Records of captive Asian elephants (Elephas maximus) were used to derive parameters of the von Bertalanffy function for growth in height, body weight and circumference of tusks with age. There was some evidence for a post-pubertal secondary growth spurt in both male and female elephants. Domestic elephants which were born in captivity or captured at a young age also showed a reduced growth in height in both the sexes and in body weight in males compared to wild elephants. Aspects of allometric growth such as height-body weight relationship are examined. The height was twice the circumference of front foot throughout the life span, indicating an isometric relationship.

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

The Asian elephant's 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 elephant's optimal foraging strategy.

Stable carbon isotope ratios in Asian elephant collagen: implications for dietary studies

Stable carbon isotope ratios in bone collagen have been used in a variety of dietary studies in modern and fossil animals, including humans. Inherent in the stable isotope technique is the assumption that the isotopic signature is a reflection of the diet and is persistent in collagen because this is a relatively inert protein. Carbon isotope analyses of bones from a southern Indian population of Asian elephant (Elephas maximus), a long-lived mammal that alternates seasonally between a predominantly C3 (browse) and C4 (grass) plant diet, showed two patterns that have important implications for dietary interpretation based on isotopic studies. Relative to the quantity of the two plant types consumed on average, the ?13C signal in collagen indicated that more carbon was incorporated from C3 plants, possibly due to their higher protein contribution. There was a much greater variance in ?13C values of collagen in sub-adult (range -10.5� to-22.7�, variance=14.51) compared to adult animals (range -16.0� to -20.3�, variance=1.85) pointing to high collagen turnover rates and non-persistent isotopic signatures in younger, growing animals. It thus seems important to correct for any significant relative differences in nutritive value of food types and also consider the age of an animal before drawing definite conclusions about its diet from isotope ratios.

Assessment of estrus cyclicity in the Asian elephant (Elephas maximus) by measurement of fecal progesterone metabolite 5 alpha-P-3OH, using a non-invasive assay

Reproductive management of the Asian elephant (Elephas maximus) is important for its conservation. To monitor its estrous cyclicity, we earlier used an indirect ELISA to show that levels of fecal progesterone (P(4))-metabolite (allopregnanolone: 5 alpha-P-3OH) in semi-captive females sampled randomly positively correlated with serum P(4) levels [12]. In this longitudinal study (51 weeks), we measured levels of fecal 5 alpha-P-3OH and serum P(4) in seven semi-captive female elephants. Females exhibited three types of hormonal profiles. Four females showed cyclical patterns of fecal 5 alpha-P-3OH and serum P(4) typical of normal estrous cycles, two showed acyclic pattern while one showed high values indicative of a pregnant animal. Values for anestrous or follicular phases were <= 0.3 mu g g(-1), (5 alpha-P-3OH) and <= 0.3 ng mL(-1) (P(4)); for luteal phase 0.32-11.09 mu g g(-1) (5 alpha-P-3OH) and 0.32-1.48 ng mL(-1) (P(4)); for pregnancy 1.41-7.38 mu g g(-1) (5 alpha-P-3OH) and 0.39-1.6 ng mL(-1) (R(4)). A positive correlation (t = 8.8, p < 0.01, n = 321) between levels of fecal 5 alpha-P-3OH and serum P4 was observed. A random sample of 30 free-ranging female elephants showed fecal 5 alpha-P-3OH values of 0.06-23.4 mu g g(-1), indicating them to be in different stages of estrous cyclicity. This study is the first to assess the reproductive phases of female Asian elephants based on the correlative-patterns of both the fecal 5 alpha-P-3OH and serum P(4) values over multiple estrous cycles. This has a potential application in the reproductive management and conservation of Asian elephants. (C) 2011 Elsevier Inc. All rights reserved.

A landscape-level assessment of Asian elephant habitat, its population and elephant-human conflict in the Anamalai hill ranges of southern Western Ghats, India

Spatial information at the landscape scale is extremely important for conservation planning, especially in the case of long-ranging vertebrates. The biodiversity-rich Anamalai hill ranges in the Western Ghats of southern India hold a viable population for the long-term conservation of the Asian elephant. Through rapid but extensive field surveys we mapped elephant habitat, corridors, vegetation and land-use patterns, estimated the elephant population density and structure, and assessed elephant-human conflict across this landscape. GIS and remote sensing analyses indicate that elephants are distributed among three blocks over a total area of about 4600 km(2). Approximately 92% remains contiguous because of four corridors; however, under 4000 km2 of this area may be effectively used by elephants. Nine landscape elements were identified, including five natural vegetation types, of which tropical moist deciduous forest is dominant. Population density assessed through the dung count method using line transects covering 275 km of walk across the effective elephant habitat of the landscape yielded a mean density of 1.1 (95% Cl = 0.99-1.2) elephant/km(2). Population structure from direct sighting of elephants showed that adult male elephants constitute just 2.9% and adult females 42.3% of the population with the rest being subadults (27.4%), juveniles (16%) and calves (11.4%). Sex ratios show an increasing skew toward females from juvenile (1:1.8) to sub-adult (1:2.4) and adult (1:14.7) indicating higher mortality of sub-adult and adult males that is most likely due to historical poaching for ivory. A rapid questionnaire survey and secondary data on elephant-human conflict from forest department records reveals that villages in and around the forest divisions on the eastern side of landscape experience higher levels of elephant-human conflict than those on the western side; this seems to relate to a greater degree of habitat fragmentation and percentage farmers cultivating annual crops in the east. We provide several recommendations that could help maintain population viability and reduce elephant-human conflict of the Anamalai elephant landscape. (C) 2013 Deutsche Gesellschaft far Saugetierkunde. Published by Elsevier GmbH. All rights reserved.

Community-managed forests and wildlife-friendly agriculture play a subsidiary but not substitutive role to protected areas for the endangered Asian elephant

Global conservation policy is increasingly debating the feasibility of reconciling wildlife conservation and human resource requirements in land uses outside protected areas (PAs). However, there are few quantitative assessments of whether or to what extent these `wildlife-friendly' land uses fulfill a fundamental function of PAs-to separate biodiversity from anthropogenic threats. We distinguish the role of wildlife-friendly land uses as being (a) subsidiary, whereby they augment PAs with secondary habitat, or (b) substitutive, wherein they provide comparable habitat to PAs. We tested our hypotheses by investigating the influence of land use and human presence on space-use intensity of the endangered Asian elephant (Elephas maximus) in a fragmented landscape comprising PAs and wildlife-friendly land uses. We applied multistate occupancy models to spatial data on elephant occurrence to estimate and model the overall probability of elephants using a site, and the conditional probability of high-intensity use given that elephants use a site. The probability of elephants using a site regardless of intensity did not vary between PAs and wildlife-friendly land uses. However, high-intensity use declined with distance to PM, and this effect was accentuated by an increase in village density. Therefore, while wildlife-friendly land uses did play a subsidiary conservation role, their potential to substitute for PAs was offset by a strong human presence. Our findings demonstrate the need to evaluate the role of wildlife-friendly land uses in landscape-scale conservation; for species that have conflicting resource requirements with people, PAs are likely to provide crucial refuge from growing anthropogenic threats. (C) 2014 Elsevier Ltd. All rights reserved.

Interplay of male traits, male mating strategies and female mate choice in the Asian elephant, Elephas maximus

Elaborate male traits with no apparent adaptive value may have evolved through female mate discrimination. Tusks are an elaborate male-only trait in the Asian elephant that could potentially influence female mate choice. We examined the effect of male body size, tusk possession and musth status on female mate choice in an Asian elephant population. Large/musth males received positive responses from oestrous females towards courtship significantly more often than did small/non-musth males. Young, tusked non-musth males attempted courtship significantly more often than their tuskless peers, and received more positive responses (though statistically insignificant) than did tuskless males. A positive response did not necessarily translate into mating because of mate-guarding by a dominant male. Female elephants appear to choose mates based primarily on traits such as musth that signal direct fertility benefits through increased sperm received than for traits such as tusks that may signal only indirect fitness benefits.

The Asian elephant is amongst the top three frugivores of two tree species with easily edible fruit

Large animal species are prone to local extirpation, but ecologists cannot yet predict how the loss of megaherbivores affects ecosystem processes such as seed dispersal. Few studies have compared the quantity and quality of seed dispersal by megaherbivores versus alternative frugivores in the wild, particularly for plant species with fruit easily consumed by many frugivorous species. In a disturbed tropical moist forest in India, we examine whether megaherbivores are a major frugivore of two tree species with easily edible, mammal-dispersed fruit. We quantify the relative fruit removal rates of Artocarpus chaplasha and Careya arborea, by the Asian elephant (Elephas maximus) and alternative dispersers. Through focal watches and camera trapping, we found the elephant to be amongst the top three frugivores for each tree species. Furthermore, seed transects under A. chaplasha show that arboreal frugivores discard seeds only a short distance from the parental tree, underscoring the elephant's role as a long-distance disperser. Our data provide unprecedented support for an old notion: megaherbivores may be key dispersers for a broad set of mammal-dispersed fruiting species, and not just fruit inaccessible to smaller frugivores. As such, the elephant may be particularly important for the functional ecology of the disturbed forests it still inhabits across tropical Asia.

Comparative sequence analyses of genome and transcriptome reveal novel transcripts and variants in the Asian elephant Elephas maximus

The Asian elephant Elephas maximus and the African elephant Loxodonta africana that diverged 5-7 million years ago exhibit differences in their physiology, behaviour and morphology. A comparative genomics approach would be useful and necessary for evolutionary and functional genetic studies of elephants. We performed sequencing of E. maximus and map to L. africana at similar to 15X coverage. Through comparative sequence analyses, we have identified Asian elephant specific homozygous, non-synonymous single nucleotide variants (SNVs) that map to 1514 protein coding genes, many of which are involved in olfaction. We also present the first report of a high-coverage transcriptome sequence in E. maximus from peripheral blood lymphocytes. We have identified 103 novel protein coding transcripts and 66-long non-coding (lnc)RNAs. We also report the presence of 181 protein domains unique to elephants when compared to other Afrotheria species. Each of these findings can be further investigated to gain a better understanding of functional differences unique to elephant species, as well as those unique to elephantids in comparison with other mammals. This work therefore provides a valuable resource to explore the immense research potential of comparative analyses of transcriptome and genome sequences in the Asian elephant.

How much Dillenia indica seed predation occurs from Asian elephant dung?

Elephants are thought to be effective seed dispersers, but research on whether elephant dung effectively protects seeds from seed predation is lacking. Quantifying rates of seed predation from elephant dung will facilitate comparisons between elephants and alternative dispersers, helping us understand the functional role of megaherbivores in ecosystems. We conducted an experiment to quantify the predation of Dillenia indica seeds from elephant dung in Buxa Reserve, India from December 2012 to April 2013. Using dung boluses from the same dung pile, we compared the number of seeds in boluses that are a) opened immediately upon detection (control boluses), b) made available only to small seed predators (<= 3 mm wide) for 1-4 months, and c) made available to all seed predators and secondary dispersers for 1-4 months. Using a model built on this experiment, we estimated that seed predation by small seed predators (most likely ants and termites) destroys between 82.9% and 96.4% of seeds in elephant dung between the time of defecation and the median germination date for D. indica. Exposure to larger seed predators and secondary dispersers did not lead to a significant additional reduction in the number of seeds per dung bolus. Our findings suggest that post-dispersal seed predation by small insects (<3 mm) substantially reduces but does not eliminate the success of elephants as dispersers of D. indica in a tropical moist forest habitat. (C) 2015 Elsevier Masson SAS. All rights reserved.

The net benefit of saving the Asian elephant: a policy and contingent valuation study

Reports results from a contingent valuation (CV) survey of willingness to pay (WTP) for the conservation of the Asian elephant of a sample of urban residents living in three selected housing schemes in Colombo, the capital of Sri Lanka. Face-to-face surveys were conducted using an interview schedule (IS). A non-linear logit regression model is used to analyse the respondents' responses for the payment principle questions and to identify the factors that influence their responses. We investigate whether urban residents' WTP for the conservation of elephants is sufficient to compensate farmers for the damage caused by elephants. We find that the beneficiaries (the urban residents) could compensate losers (the fanners in the areas affected by human-elephant conflict, HEC) and be better off than in the absence of elephants in Sri Lanka. Therefore, there is a strong economic case for the conservation of the wild elephant population in Sri Lanka. However, we have insufficient data to determine the optimal level of this elephant population in the Kaldor-Hicks sense. Nevertheless, the current population of elephant in Sri Lanka is Kaldor-Hicks preferable to having none. (C) 2003 Elsevier B.V. All rights reserved.

Vocalizations of wild Asian elephants (Elephas maximus): Structural classification and social context

Elephants use vocalizations for both long and short distance communication. Whereas the acoustic repertoire of the African elephant (Loxodonta africana) has been extensively studied in its savannah habitat, very little is known about the structure and social context of the vocalizations of the Asian elephant (Elephas maximus), which is mostly found in forests. In this study, the vocal repertoire of wild Asian elephants in southern India was examined. The calls could be classified into four mutually exclusive categories, namely, trumpets, chirps, roars, and rumbles, based on quantitative analyses of their spectral and temporal features. One of the call types, the rumble, exhibited high structural diversity, particularly in the direction and extent of frequency modulation of calls. Juveniles produced three of the four call types, including trumpets, roars, and rumbles, in the context of play and distress. Adults produced trumpets and roars in the context of disturbance, aggression, and play. Chirps were typically produced in situations of confusion and alarm. Rumbles were used for contact calling within and among herds, by matriarchs to assemble the herd, in close-range social interactions, and during disturbance and aggression. Spectral and temporal features of the four call types were similar between Asian and African elephants.