All-Male Groups in Asian Elephants: A Novel, Adaptive Social Strategy in Increasingly Anthropogenic Landscapes of Southern India.
ABSTRACT: Male Asian elephants are known to adopt a high-risk high-gain foraging strategy by venturing into agricultural areas and feeding on nutritious crops in order to improve their reproductive fitness. We hypothesised that the high risks to survival posed by increasingly urbanising and often unpredictable production landscapes may necessitate the emergence of behavioural strategies that allow male elephants to persist in such landscapes. Using 1445 photographic records of 248 uniquely identified male Asian elephants over a 23-month period, we show that male Asian elephants display striking emergent behaviour, particularly the formation of stable, long-term all-male groups, typically in non-forested or human-modified and highly fragmented areas. They remained solitary or associated in mixed-sex groups, however, within forested habitats. These novel, large all-male associations, may constitute a unique life history strategy for male elephants in the high-risk but resource-rich production landscapes of southern India. This may be especially true for the adolescent males, which seemed to effectively improve their body condition by increasingly exploiting anthropogenic resources when in all-male groups. This observation further supports our hypothesis that such emergent behaviours are likely to constitute an adaptive strategy for male Asian elephants that may be forced to increasingly confront anthropogenically intrusive environments.
Project description:BACKGROUND: A dearth in understanding the behavior of Asian elephants (Elephas maximus) at the scale of populations and individuals has left important management issues, particularly related to human-elephant conflict (HEC), unresolved. Evaluation of differences in behavior and decision-making among individual elephants across groups in response to changing local ecological settings is essential to fill this gap in knowledge and to improve our approaches towards the management and conservation of elephants. METHODOLOGY/PRINCIPAL FINDINGS: We hypothesized certain behavioral decisions that would be made by Asian elephants as reflected in their residence time and movement rates, time-activity budgets, social interactions and group dynamics in response to resource availability and human disturbance in their habitat. This study is based on 200 h of behavioral observations on 60 individually identified elephants and a 184-km(2) grid-based survey of their natural and anthropogenic habitats within and outside the Bannerghatta National Park, southern India during the dry season. At a general population level, the behavioral decisions appeared to be guided by the gender, age and group-type of the elephants. At the individual level, the observed variation could be explained only by the idiosyncratic behaviors of individuals and that of their associating conspecific individuals. Recursive partitioning classification trees for residence time of individual elephants indicated that the primary decisions were taken by individuals, independently of their above-mentioned biological and ecological attributes. CONCLUSIONS/SIGNIFICANCE: Decision-making by Asian elephants thus appears to be determined at two levels, that of the population and, more importantly, the individual. Models based on decision-making by individual elephants have the potential to predict conflict in fragmented landscapes that, in turn, could aid in mitigating HEC. Thus, we must target individuals, in addition to populations, in our efforts to manage and conserve this threatened species, particularly in human-dominated landscapes.
Project description:Loss of forest cover, rise in human populations and fragmentation of habitats leads to decline in biodiversity and extinction of large mammals globally. Elephants, being the largest of terrestrial mammals, symbolize global conservation programs and co-occur with humans within multiple-use landscapes of Asia and Africa. Within such shared landscapes, poaching, habitat loss and extent of human-elephant conflicts (HEC) affect survival and conservation of elephants. HEC are severe in South Asia with increasing attacks on humans, crop depredation and property damage. Such incidents reduce societal tolerance towards elephants and increase the risk of retaliation by local communities. We analyzed a 2-year dataset on crop depredation by Asian elephants (N = 380) events in North Bengal (eastern India). We also explored the effect of landscape, anthropogenic factors (area of forest, agriculture, distance to protected area, area of human settlements, riverine patches and human density) on the spatial occurrence of such incidents.Crop depredation showed a distinct nocturnal pattern (22.00-06:00) and majority of the incidents were recorded in the monsoon and post-monsoon seasons. Results of our spatial analysis suggest that crop depredation increased with an increase in the area of forest patches, agriculture, presence of riverine patches and human density. Probability of crop depredation further increased with decreasing distance from protected areas. Villages within 1.5 km of a forest patch were most affected. Crop raiding incidents suggest a deviation from the "high-risk high-gain male biased" foraging behavior and involved proportionately more mixed groups (57%) than lone bulls (43%). Demographic data suggest that mixed groups comprised an average of 23 individuals with adult and sub adult females, bulls and calves. Crop depredation and fatal elephant attacks on humans were spatially clustered with eastern, central and western parts of North Bengal identified as hotspots of HEC. Our results will help to prioritize mitigation measures such as prohibition of alcohol production within villages, improving condition of riverine patches, changing crop composition, fencing agriculture fields, implement early warning systems around protected areas and training local people on how to prevent conflicts.
Project description:To elucidate the history of living and extinct elephantids, we generated 39,763 bp of aligned nuclear DNA sequence across 375 loci for African savanna elephant, African forest elephant, Asian elephant, the extinct American mastodon, and the woolly mammoth. Our data establish that the Asian elephant is the closest living relative of the extinct mammoth in the nuclear genome, extending previous findings from mitochondrial DNA analyses. We also find that savanna and forest elephants, which some have argued are the same species, are as or more divergent in the nuclear genome as mammoths and Asian elephants, which are considered to be distinct genera, thus resolving a long-standing debate about the appropriate taxonomic classification of the African elephants. Finally, we document a much larger effective population size in forest elephants compared with the other elephantid taxa, likely reflecting species differences in ancient geographic structure and range and differences in life history traits such as variance in male reproductive success.
Project description:The accelerating expansion of human populations and associated economic activity across the globe have made maintaining large, intact natural areas increasingly challenging. The difficulty of preserving large intact landscapes in the presence of growing human populations has led to a growing emphasis on landscape approaches to biodiversity conservation with a complementary strategy focused on improving conservation in human-modified landscapes. This, in turn, is leading to intense debate about the effectiveness of biodiversity conservation in human-modified landscapes and approaches to better support biodiversity in those landscapes. Here, we compared butterfly abundance, alpha richness, and beta diversity in human-modified landscapes (urban, sugarcane) and natural, forested areas to assess the conservation value of human-modified landscapes within the Wet Tropics bioregion of Australia. We used fruit-baited traps to sample butterflies and analyzed abundance and species richness in respective land uses over a one-year period. We also evaluated turnover and spatial variance components of beta diversity to determine the extent of change in temporal and spatial variation in community composition. Forests supported the largest numbers of butterflies, but were lowest in each, alpha species richness, beta turnover, and the spatial beta diversity. Sugarcane supported higher species richness, demonstrating the potential for conservation at local scales in human-modified landscapes. In contrast, beta diversity was highest in urban areas, likely driven by spatial and temporal variation in plant composition within the urban landscapes. Thus, while improving conservation on human-modified landscapes may improve local alpha richness, conserving variation in natural vegetation is critical for maintaining high beta diversity.
Project description:Personality, i.e. consistent between-individual differences in behaviour, has been documented in many species. Yet little is known about how males and females of long-lived, highly social species differ in their measures of personality structure. We investigated sex differences in the mean, variance, and covariance of three previously reported personality traits (Attentiveness, Sociability, Aggressiveness) in 150 female and 107 male Asian elephants (Elephas maximus) from a semi-captive population in Myanmar. These three personality traits were obtained by performing exploratory factor analysis on 28 behavioural items that had been rated by experienced elephant handlers. We found that males scored significantly higher on Aggressiveness and tended to score lower on Sociability than females. However, no sex difference was found in the mean scores of Attentiveness. Variances for the three personality traits did not differ between the sexes, suggesting that male and female elephants share the same range of personality variation. Likewise, trait covariances were similar between the sexes. While both sexes show complex sociality in the wild, female Asian elephants typically live in highly social family units, whereas male elephants' social bonds are weaker. Males usually form dominance ranks by aggressive interactions, especially during musth. Our results on a large sample of individuals living in their natural environment are thus in agreement with elephant life-histories and parallel the findings of sex differences in other long-lived highly social species with similar life-histories.
Project description:Salivary immunoglobulin A (sIgA) has been proposed as a potential indicator of welfare for various species, including Asian elephants, and may be related to adrenal cortisol responses. This study aimed to distinguish circadian rhythm effects on sIgA in male and female Asian elephants and compare patterns to those of salivary cortisol, information that could potentially have welfare implications. Subjects were captive elephants at an elephant camp in Chiang Mai province, Thailand (n = 5 males, 5 females). Salivette® kits were used to collect saliva from each elephant every 4 h from 06:00 to 22:00 h for 3 consecutive days (n = 15 samples/elephant). Enzyme immunoassays were used to quantify concentrations of IgA and cortisol in unextracted saliva. Circadian rhythm patterns were determined using a generalized least-squares method. Both sIgA and cortisol followed a circadian rhythm, although the patterns differed. sIgA displayed a daily quartic trend, whereas cortisol concentrations demonstrated a decreasing linear trend in concentrations throughout the day. There was no clear relationship between patterns of sIgA and salivary cortisol, implying that mechanisms of control and secretion differ. Results demonstrate for the first time that circadian rhythms affect sIgA, and concentrations follow a daily quartic pattern in Asian elephants, so standardizing time of collection is necessary.
Project description: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.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.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.
Project description:Resting behaviors are an essential component of animal welfare but have received little attention in zoological research. African savanna elephant (Loxodonta africana) and Asian elephant (Elephas maximus) rest includes recumbent postures, but no large-scale investigation of African and Asian zoo elephant recumbence has been previously conducted. We used anklets equipped with accelerometers to measure recumbence in 72 adult female African (n = 44) and Asian (n = 28) elephants housed in 40 North American zoos. We collected 344 days of data and determined associations between recumbence and social, housing, management, and demographic factors. African elephants were recumbent less (2.1 hours/day, S.D. = 1.1) than Asian elephants (3.2 hours/day, S.D. = 1.5; P < 0.001). Nearly one-third of elephants were non-recumbent on at least one night, suggesting this is a common behavior. Multi-variable regression models for each species showed that substrate, space, and social variables had the strongest associations with recumbence. In the African model, elephants who spent any amount of time housed on all-hard substrate were recumbent 0.6 hours less per day than those who were never on all-hard substrate, and elephants who experienced an additional acre of outdoor space at night increased their recumbence by 0.48 hours per day. In the Asian model, elephants who spent any amount of time housed on all-soft substrate were recumbent 1.1 hours more per day more than those who were never on all-soft substrate, and elephants who spent any amount of time housed alone were recumbent 0.77 hours more per day than elephants who were never housed alone. Our results draw attention to the significant interspecific difference in the amount of recumbent rest and in the factors affecting recumbence; however, in both species, the influence of flooring substrate is notably important to recumbent rest, and by extension, zoo elephant welfare.
Project description: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.
Project description:Elephant endotheliotropic herpesviruses (EEHVs) can cause fatal hemorrhagic disease in juvenile Asian elephants (Elphas maximus); however, sporadic shedding of virus in trunk washes collected from healthy elephants also has been detected. Data regarding the relationship of viral loads in blood compared with trunk washes are lacking, and questions about whether elephants can undergo multiple infections with EEHVs have not been addressed previously. Real-time quantitative polymerase chain reaction was used to determine the kinetics of EEHV1 loads, and genotypic analysis was performed on EEHV1 DNA detected in various fluid samples obtained from five Asian elephants that survived detectable EEHV1 DNAemia on at least two separate occasions. In three elephants displaying clinical signs of illness, preclinical EEHV1 DNAemia was detectable, and peak whole-blood viral loads occurred 3-8 days after the onset of clinical signs. In two elephants with EEHV1 DNAemia that persisted for 7-21 days, no clinical signs of illness were observed. Detection of EEHV1 DNA in trunk washes peaked approximately 21 days after DNAemia, and viral genotypes detected during DNAemia matched those detected in subsequent trunk washes from the same elephant. In each of the five elephants, two distinct EEHV1 genotypes were identified in whole blood and trunk washes at different time points. In each case, these genotypes represented both an EEHV1A and an EEHV1B subtype. These data suggest that knowledge of viral loads could be useful for the management of elephants before or during clinical illness. Furthermore, sequential infection with both EEHV1 subtypes occurs in Asian elephants, suggesting that they do not elicit cross-protective sterilizing immunity. These data will be useful to individuals involved in the husbandry and clinical care of Asian elephants.