Population genetic structure and conservation of Asian elephants (Elephas maximus) across India

This study examines the population genetic structure of Asian elephants (Elephas maximus) across India, which harbours over half the world's population of this endangered species. Mitochondrial DNA control region sequences and allele frequencies at six nuclear DNA microsatellite markers obtained from the dung of free-ranging elephants reveal low mtDNA and typical microsatellite diversity. Both known divergent clades of mtDNA haplotypes in the Asian elephant are present in India, with southern and central India exhibiting exclusively the β clade of Fernando et al. (2000), northern India exhibiting exclusively the α clade and northeastern India exhibiting both, but predominantly the α clade. A nested clade analysis revealed isolation by distance as the principal mechanism responsible for the observed haplotype distributions within the α and β clades. Analyses of molecular variance and pairwise population FST tests based on both mitochondrial and microsatellite DNA suggest that northern-northeastern India, central India, Nilgiris (in southern India) and Anamalai-Periyar (in southern India) are four demographically autonomous population units and should be managed separately. In addition, evidence for female philopatry, male-mediated gene flow and two possible historical biogeographical barriers is described.

The Asian elephants Elephas maximus of Cat Tien National Park, Vietnam: status and conservation of a vanishing population

This study updates the status and conservation of the Endangered Asian elephant Elephas maximus in Cat Tien National Park, Vietnam. Line transect indirect surveys, block surveys for elephant signs, village surveys of elephant-human conflict incidents, guard-post surveys for records of sightings, and surveys of elephant food plants were undertaken during the dry and wet seasons of 2001. A minimum of 11 elephants and a maximum of 15-17 elephants was estimated for c. 500 km2 of the Park and its vicinity. The elephants are largely confined to the southern boundary of the Park and make extensive use of the adjoining La Nga State Forest Enterprises. During the dry season the elephants depend on at least 26 species of wild and cultivated plants, chiefly the fruits of cashew. Most of the villages surveyed reported some elephant-human conflict. Two adult male elephants seem to cover a large area to raid crops, whereas the family groups restrict themselves to a few villages; overall, the conflict is not serious. Since 2001 there have been no reports of any deaths or births of elephants in the Park. We make recommendations for habitat protection and management, increasing the viability of the small population, reducing elephant-human conflicts, and improving the chances of survival of the declining elephants of this Park. The Government has now approved an Action Plan for Urgent Conservation Areas in Vietnam that calls for the establishment of three elephant conservation areas in the country, including Cat Tien National Park.

DNA analysis indicates that Asian elephants are native to Borneo and are therefore a high priority for conservation

The origin of Borneo's elephants is controversial. Two competing hypotheses argue that they are either indigenous, tracing back to the Pleistocene, or were introduced, descending from elephants imported in the 16th-18th centuries. Taxonomically, they have either been classified as a unique subspecies or placed under the Indian or Sumatran subspecies. If shown to be a unique indigenous population, this would extend the natural species range of the Asian elephant by 1300 km, and therefore Borneo elephants would have much greater conservation importance than if they were a feral population. We compared DNA of Borneo elephants to that of elephants from across the range of the Asian elephant, using a fragment of mitochondrial DNA, including part of the hypervariable d-loop, and five autosomal microsatellite loci. We find that Borneo's elephants are genetically distinct, with molecular divergence indicative of a Pleistocene colonisation of Borneo and subsequent isolation. We reject the hypothesis that Borneo's elephants were introduced. The genetic divergence of Borneo elephants warrants their recognition as a separate evolutionary significant unit. Thus, interbreeding Borneo elephants with those from other populations would be contraindicated in ex situ conservation, and their genetic distinctiveness makes them one of the highest priority populations for Asian elephant conservation.

Dung as a potential medium for inter-sexual chemical signaling in Asian elephants (Elephas maximus)

Chemical signaling is a prominent mode of male-female communication among elephants, especially during their sexually active periods. Studies on the Asian elephant in zoos have shown the significance of a urinary pheromone (Z7-12:Ac) in conveying the reproductive status of a female toward the opposite sex. We investigated the additional possibility of an inter-sexual chemical signal being conveyed through dung. Sixteen semi-captive adult male elephants were presented with dung samples of three female elephants in different reproductive phases. Each male was tested in 3 separate trials, within an interval of 1-3 days. The trials followed a double-blind pattern as the male and female elephants used in the trials were strangers, and the observer was not aware of the reproductive status of females during the period of bioassays. Males responded preferentially (P < 0.005), in terms of higher frequency of sniff, check and place behavior toward the dung of females close to pre-ovulatory period (follicular-phase) as compared to those in post-ovulatory period (luteal-phase). The response toward the follicular phase samples declined over repeated trials though was still significantly higher than the corresponding response toward the non-ovulatory phase in each of the trials performed. This is the first study to show that male Asian elephants were able to distinguish the reproductive phase of the female by possibly detecting a pre-ovulatory pheromone released in dung. (C) 2012 Elsevier B.V. All rights reserved.

Modeling harvest rates and numbers from age and sex ratios: A demonstration for elephant populations

Illegal harvest rates of wildlife populations are often unknown or difficult to estimate from field data due to under-reporting or incomplete detection of carcasses. This is especially true for elephants that are killed for ivory or in conflicts with people. We describe a method to infer harvest rates from coarse field data of three population parameters, namely, adult female to male ratio, male old-adult to young-adult ratio, and proportion of adult males in the population using Jensen's (2000) 2-sex, density-dependent Leslie matrix model. The specific combination of male and female harvest rates and numbers can be determined from the history of harvest and estimate of population size. We applied this technique to two populations of elephants for which data on age structure and records of mortality were available-a forest-dwelling population of the Asian elephant (at Nagarahole, India) and an African savannah elephant population (at Samburu, Kenya) that had experienced male-biased harvest regimes over 2-3 decades. For the Nagarahole population, the recorded numbers of male and female elephants killed illegally during 1981-2000 were 64% and 88% of the values predicted by the model, respectively, implying some non-detection or incomplete reporting while for the Samburu population the recorded and modeled numbers of harvest during 1990-1999 closely matched. This technique, applicable to any animal population following logistic growth model, can be especially useful for inferring illegal harvest numbers of forest elephants in Africa and Asia.

The role of tusks, musth and body size in male-male competition among Asian elephants, Elephas maximus

The evolution of sexually dimorphic, elaborate male traits that are seemingly maladaptive may be driven by sexual selection (male-male competition and or female mate choice). Tusk possession in the Asian elephant is sexually dimorphic and exaggerated but its role in male-male competition has not yet been determined. We examined the role of the tusks in establishing dominance along with two other known male-male signals, namely, body size and musth (a temporary physiologically heightened sexual state) in an Asian elephant population in northeastern India with equal proportions of tusked and tuskless males. We observed 116 agonistic interactions with clear dominance outcomes between adult (>15 years) males during 458 field days in the dry season months of 2008-2011. A generalized linear mixed-effects model was used to predict the probability of winning as a function of body size, tusk possession and musth status relative to the opponent. A hierarchy of the three male-male signals emerged from this analysis, with musth overriding body size and body size overriding tusk possession. In this elephant population tusk possession thus plays a relatively minor role in male-male competition. An important implication of musth and body size being stronger determinants of dominance than tusk possession is that it could facilitate rapid evolution of tuskless males in the population under artificial selection against tusked individuals, which are poached for ivory. (C) 2013 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Endocrine Correlates of Musth in Free-Ranging Asian Elephants (Elephas maximus) Determined by Non-Invasive Faecal Steroid Hormone Metabolite Measurements

The occurrence of musth, a period of elevated levels of androgens and heightened sexual activity, has been well documented for the male Asian elephant (Elephas maximus). However, the relationship between androgen-dependent musth and adrenocortical function in this species is unclear. The current study is the first assessment of testicular and adrenocortical function in free-ranging male Asian elephants by measuring levels of testosterone (androgen) and cortisol (glucocorticoid - a physiological indicator of stress) metabolites in faeces. During musth, males expectedly showed significant elevation in faecal testosterone metabolite levels. Interestingly, glucocorticoid metabolite concentrations remained unchanged between musth and non-musth periods. This observation is contrary to that observed with wild and captive African elephant bulls and captive Asian bull elephants. Our results show that musth may not necessarily represent a stressful condition in free-ranging male Asian elephants.

Distribution, relative abundance, and conservation status of Asian elephants in Karnataka, southern India

Karnataka state in southern India supports a globally significant and the country's largest population of the Asian elephant Elephas maximus. A reliable map of Asian elephant distribution and measures of spatial variation in their abundance, both vital needs for conservation and management action, are unavailable not only in Karnataka, but across its global range. Here, we use various data gathered between 2000 and 2015 to map the distribution of elephants in Karnataka at the scale of the smallest forest management unit, the `beat', while also presenting data on elephant dung density for a subset of `elephant beats.' Elephants occurred in 972 out of 2855 forest beats of Karnataka. Sixty percent of these 972 beats and 55% of the forest habitat lay outside notified protected areas (PM), and included lands designated for agricultural production and human dwelling. While median elephant dung density inside protected areas was nearly thrice as much as outside, elephants routinely occurred in or used habitats outside PM where human density, land fraction under cultivation, and the interface between human-dominated areas and forests were greater. Based on our data, it is clear that India's framework for elephant conservation which legally protects the species wherever it occurs, but protects only some of its habitats while being appropriate in furthering their conservation within PM, seriously falters in situations where elephants reside in and/or seasonally use areas outside PAs. Attempts to further elephant conservation in production and dwelling areas have extracted high costs in human, elephant, material and monetary terms in Karnataka. In such settings, conservation planning exercises are necessary to determine where the needs of elephants or humans must take priority over the other, and to achieve that in a manner that is based not only on reliable scientific data but also on a process of public reasoning. (C) 2015 Elsevier Ltd. All rights reserved.

Modeling harvest rates and numbers from age and sex ratios: A demonstration for elephant populations

Illegal harvest rates of wildlife populations are often unknown or difficult to estimate from field data due to under-reporting or incomplete detection of carcasses. This is especially true for elephants that are killed for ivory or in conflicts with people. We describe a method to infer harvest rates from coarse field data of three population parameters, namely, adult female to male ratio, male old-adult to young-adult ratio, and proportion of adult males in the population using Jensen's (2000) 2-sex, density-dependent Leslie matrix model. The specific combination of male and female harvest rates and numbers can be determined from the history of harvest and estimate of population size. We applied this technique to two populations of elephants for which data on age structure and records of mortality were available-a forest-dwelling population of the Asian elephant (at Nagarahole, India) and an African savannah elephant population (at Samburu, Kenya) that had experienced male-biased harvest regimes over 2-3 decades. For the Nagarahole population, the recorded numbers of male and female elephants killed illegally during 1981-2000 were 64% and 88% of the values predicted by the model, respectively, implying some non-detection or incomplete reporting while for the Samburu population the recorded and modeled numbers of harvest during 1990-1999 closely matched. This technique, applicable to any animal population following logistic growth model, can be especially useful for inferring illegal harvest numbers of forest elephants in Africa and Asia. (C) 2013 Elsevier Ltd. All rights reserved.

Comparison of rural and urban attitudes to the conservation of Asian elephants in Sri Lanka: empirical evidence

There has been little study of economic and general attitudes towards the conservation of the Asian elephant. This paper reports and analyses results from surveys conducted in Sri Lanka of attitudes of urban dwellers and farmers towards nature conservation in general and the elephant conservation in particular. The analyses are based on urban and a rural sample. Contingent valuation techniques are used as survey instruments. Multivariate logit regression analysis is used to analyse the respondents' attitudes towards conservation of elephants. It is found that, although some variations occurred between the samples, the majority of the respondents (both rural and urban) have positive attitudes towards nature conservation in general. However, marked differences in attitudes toward elephant conservation are evident between these two samples: the majority of urban respondents were in favour of elephant conservation; rural respondents expressed a mixture of positive and negative attitudes. Overall, considerable unrecorded and as yet unutilised economic support for conservation of wild elephants exists in Sri Lanka. (C) 2002 Elsevier Science Ltd. All rights reserved.

SOCIAL STRUCTURE OF ASIAN ELEPHANTS (ELEPHAS MAXIMUS) IN SRI LANKA

Asian elephants (Elephas maximus) are critically endangered and live in fragmented populations spread across 13 countries. Yet in comparison to the African savannah elephant (Loxodonta africana), relatively little is known about the social structure of wild Asian elephants because the species is mostly found in low visibility habitat. A better understanding of Asian elephant social structure is critical to mitigate human-elephant conflicts that arise due to increasing human encroachments into elephant habitats. In this dissertation, I examined the social structure of Asian elephants at three sites: Yala, Udawalawe, and Minneriya National Parks in Sri Lanka, where the presence of large open areas and high elephant densities are conducive to behavioral observations. First, I found that the size of groups observed at georeferenced locations was affected by forage availability and distance to water, and the effects of these environmental factors on group size depended on site. Second, I discovered that while populations at different sites differed in the prevalence of weak associations among individuals, a core social structure of individuals sharing strong bonds and organized into highly independent clusters was present across sites. Finally, I showed that the core social structure preserved across sites was typically composed of adult females associating with each other and with other age-sex classes. In addition, I showed that females are social at all life stages, whereas males gradually transition from living in a group to a more solitary lifestyle. Taking into consideration these elements of Asian elephant social structure will help conservation biologists develop effective management strategies that account for both human needs and the socio-ecology of the elephants.

A novel antigen capture ELISA for the specific detection of IgG antibodies to elephant endotheliotropic herpes virus

BACKGROUND Elephants are classified as critically endangered animals by the International Union for Conservation of Species (IUCN). Elephant endotheliotropic herpesvirus (EEHV) poses a large threat to breeding programs of captive Asian elephants by causing fatal haemorrhagic disease. EEHV infection is detected by PCR in samples from both clinically ill and asymptomatic elephants with an active infection, whereas latent carriers can be distinguished exclusively via serological assays. To date, identification of latent carriers has been challenging, since there are no serological assays capable of detecting seropositive elephants. RESULTS Here we describe a novel ELISA that specifically detects EEHV antibodies circulating in Asian elephant plasma/serum. Approximately 80 % of PCR positive elephants display EEHV-specific antibodies. Monitoring three Asian elephant herds from European zoos revealed that the serostatus of elephants within a herd varied from non-detectable to high titers. The antibody titers showed typical herpes-like rise-and-fall patterns in time which occur in all seropositive animals in the herd more or less simultaneously. CONCLUSIONS This study shows that the developed ELISA is suitable to detect antibodies specific to EEHV. It allows study of EEHV seroprevalence in Asian elephants. Results confirm that EEHV prevalence among Asian elephants (whether captive-born or wild-caught) is high.

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 Mudumalai 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 gauras). 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 analysing 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 are 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 chital; 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 af the forest, indicating that road-side counts may not be appropriate for most species.

Phylogenetic resolution within the Elephantidae using fossil DNA sequence from the American mastodon (Mammut americanum) as an outgroup.

DNA was extracted from the extinct American mastodon, the extinct woolly mammoth, and the modern Asian and African elephants to test the traditional morphologically based phylogeny within Elephantidae. Phylogenetic analyses of the aligned sequences of the mitochondrial gene cytochrome b support a monophyletic Asian elephant-woolly mammoth clade when the American mastodon is used as an outgroup. Previous molecular studies were unable to resolve the relationships of the woolly mammoth, Asian elephant, and African elephant because the sequences appear to have evolved at heterogeneous rates and inappropriate outgroups were used for analysis. The results demonstrate the usefulness of fossil molecular data from appropriate sister taxa for resolving phylogenies of highly derived or early radiating lineages.

Changing abundance of elephants and willingness to pay for their conservation

This paper explores the way in which the stated willingness to pay for the conservation of Asian elephants in Sri Lanka varies with hypothetical variations in their abundance. To do that, it relies on results from a sample of residents of Colombo. The willingness to pay function is found to be unusual. It increases at an increasing rate for hypothetical reductions in the elephant population compared to its current level (a level that makes the Asian elephant endangered) and also increases at a decreasing rate for increases in this population from its current level. Rational explanations are Oven for this relationship. The relationship is, however, at odds with relationships suggested in some of the literature for total economic value as a function of the abundance of a wildlife species. It is suggested that willingness to pay for conservation of a species rationally includes a strategic element and may not always measure the total economic value of it species. Nevertheless. willingness to pay is still policy relevant in such cases. (c) 2005 Elsevier Ltd. All rights reserved.