Dissertations / Theses on the topic 'Elephant, asian'

Elephant endotheliotropic herpesviruses (EEHVs) are responsible for a highly fatal haemorrhagic disease (EEHV-HD), threatening the overall sustainability of the Asian elephant (Elephas maximus) population. The viruses were first reported in tissue sections of a fatal case of an Asian elephant in 1988. In recent years, there has been a gradual increase in our understanding of the viruses highlighted by decoding their complete genome sequences. However, lack of an animal model and a cell culture system has restricted studies on the pathogenesis of the viruses, assessing control measures, and development of a vaccine. This project therefore aimed to achieve a better understanding of the viruses’ pathogenesis studying several EEHV surviving and fatal cases, and to advise accordingly on the control measures. The results revealed coincidence of highest blood viral load with clinical signs, viruses’ shedding in trunk secretion during convalescence, complementary haematological tests to aid detection and clinical prognosis of EEHV infections and the inadequacy of famciclovir treatment to alter EEHV infections. Analysis of tissues from fatal cases detected viruses’ co-infections (EEHV-1 and 4) and widespread organ distribution with liver, heart, mesenteric lymph node, thymus, and tongue having the highest viral load, thereby highlighting implications for EEHV diagnosis and future in vitro isolation. Further, this study isolated elephant endothelial cells and PBMCs and used these established elephant cells alongside common laboratory cell lines to assess their suitability for EEHV-1A isolation. Despite indication of limited increase in EEHV-1 DNA in PBMCs and mouse embryo fibroblast supernatants, conclusive in vitro virus replication could not be demonstrated. In conclusion, the outcomes of this PhD study further advance our knowledge on pathogenesis of EEHV strains and provide new insights on their control to minimise their respective impact on the Asian elephant population.

Herpesviruses are ubiquitous and are found worldwide, most animal species can be infected with multiple herpesviruses. Some cause clinical disease and others remain symptomatic throughout life. Herpesviruses are found in both captive and wild animals including Asian elephants (Elephas maximus). Elephant Endothelioltropic Herpesvirus (EEHV) has been reported in both captive and wild Asian elephants, with a number of cases being reported in North America, Europe and Asia. It has been suggested that EEHV is associated with haemorrhagic disease, which has been attributed to a number of Asian elephant deaths, affecting mostly juveniles and calves. Clinical signs can vary from weight loss, lethargy, depression, cyanosis of the tongue and sudden death. Molecular testing using qPCR has enabled the detection of individual variants of EEHV, this thesis investigates the EEHV1 variant. EEHV1 has been highlighted as the variant that is more frequently associated with deaths. This thesis includes five studies investigating different aspects of EEHV. Including, the relationship between pregnancy and EEHV viral shedding, the use of an amended human protocol for culturing endothelial cells, EEHV tissue tropism, a potential genetic or familial link between EEHV associated deaths and the detection of potential co-pathogens. The main findings from this thesis include: 1) the use of a longitudinal study investigating a potential link between the physiological stress of pregnancy and EEHV viral shedding. This study suggested there was no link between pregnancy and EEHV viral shedding however other stressors may be involved. 2) Using an amended human umbilical vein endothelial cell protocol, the culture of Asian elephant endothelial cells was successful. The cells from this study may be used in subsequent drug testing and vaccine development. 3) Quantitative PCR was used to determine EEHV1 tropism in tissues from two deaths associated with the virus. Tropism appeared to be for the heart and liver. 4) This thesis provides results from a preliminary study into a potential link between EEHV associated deaths. The data from an Asian elephant genogram shows there is the possibility of a genetic or familial link, which requires further investigation. 5) A number of tissues from deaths associated with EEHV and or death from other causes were investigated for the presence of potential co-pathogens, including the presence of encephalomyocarditis virus (EMCV), using microarray technology. The results indicated there were no co-pathogens present in the tissues. This thesis adds to the current published data, and includes the first known preliminary study investigating a potential genetic link between elephant deaths due to EEHV.

This thesis examined the effectiveness of DNA fingerprinting analysis for paternity ascertainment and the establishment of relatedness of captive Asian elephants (Elephas maximas). Eighteen Asian elephants from three North .American zoos were examined. Thirteen of these elephants were wild caught. Relationships between these elephants and the remaining elephants born in captivity were known.

Elephant Endotheliotropic Herpesvirus (EEHV) infection is of conservation concern to endangered Asian elephants (Elephas maximus), causing severe, acute, often fatal haemorrhagic disease in young elephants. This thesis investigates the epidemiological status of clinical and subclinical EEHV infection in Thailand, through a retrospective, a cross-sectional, and a longitudinal study in captive elephants. Novel diagnostic tools were developed and recommendations discussed, with a particular focus on practical recommendations to manage EEHV infection in Thailand, where logistical and management constraints can hamper disease diagnosis and timely treatment. The retrospective study revealed at least 18 EEHV clinical cases in Thailand (2006–2014), with only two cases surviving following intensive treatment. Viral samples from each case were not identical based on multiple gene analysis; suggesting the disease is likely to be sporadic. The findings also suggest that EEHV1A and EEHV1B are likely endogenous pathogens in Asian elephants. A novel SYBR green I-based real-time PCR assay was developed to identify subclinical infection, diagnose early infection, and monitor disease progression. This technique provided sufficient data, with appropriate detection limits to differentiate at least three types of EEHV (EEHV1A, 1B, and 4). Applying this technology to the cross-sectional study, conducted between 2013–2015, revealed a 5.5% prevalence of asymptomatic EEHV1 infection during this study period in captive Asian elephants in Thailand. Cross-sectional analysis did not find an association between EEHV infection and sex, location, or contact history with other EEHV-positive elephants; however, EEHV was more likely to be detected in juveniles than other age classes (OR = 4.46; 95%CI: 1.60–12.45; p = 0.05). The study also monitored EEHV shedding patterns longitudinally in an EEHV-positive herd. Frequency of EEHV detection varied within and between individuals, but was significantly higher in elephants which had survived previous clinical EEHV infection (OR = 4.85; 95%CI: 0.88–26.74; p = 0.05). Concurrent monitoring of faecal corticosterone metabolites demonstrated that EEHV activated and reactivated spontaneously and shed sporadically, despite lack of obvious stress. This thesis describes the first intensive EEHV study in an Asian elephant range country. The thesis assesses the novel real-time PCR protocol and current diagnostic tools, and recommends practical management and disease preparedness strategies to minimise the impacts of EEHV on wild and captive populations, both locally and internationally.

Relatively little is known about the vocal repertoire of Asian elephants (Elephas maximus), and a categorization of basic call types and modifications of these call types by quantitative acoustic parameters is needed to examine acoustic variability within and among call types, to examine individuality, to determine communicative function of calls via playback, to compare species and populations, and to develop rigorous call recognition algorithms for monitoring populations. This study defines an acoustic repertoire of Asian elephants based on acoustic parameters, compares repertoire usage among groups and individuals, and validates structural distinction among call types through comparison of manual and automated classification methods. Recordings were made of captive elephants at the Oregon Zoo in Portland, OR, USA, and of domesticated elephants in Thailand. Acoustic and behavioral data were collected in a variety of social contexts and environmental noise conditions. Calls were classified using perceptual aural cues plus visual inspection of spectrograms, then acoustic features were measured, then automated classification was run. The final repertoire was defined by six basic call types (Bark, Roar, Rumble, Bark, Squeal, Squeal, and Trumpet), five call combinations and modifications with these basic calls forming their constituent parts (Roar-Rumble, Squeal-Squeak, Squeak train, Squeak-Bark, and Trumpet-Roar), and the Blow. Given the consistency of classifications results for calls from geographically and socially disparate subject groups, it seems possible that automated call detection algorithms could be developed for acoustic monitoring of Asian elephants.

Behavioral evidence suggests that Asian elephants strongly rely on their sense of smell in a variety of contexts including foraging and social communication. Using a food-rewarded two-alternative operant conditioning procedure, three female Asian elephants were tested on their olfactory discrimination ability with 1-aliphatic alcohols, n-aldehydes, 2-ketones, n-carboxylic acids and with a set of twelve enantiomeric odor pairs. When presented with pairs of structurally related aliphatic odorants, the discrimination performance of the elephants increased with decreasing structural similarity of the odorants. Nevertheless, the animals successfully discriminated between all aliphatic odorants even when these only differed by one carbon atom. The elephants were also able to discriminate between all twelve enantiomeric odor pairs tested. Additionally, the elephants showed an excellent long-term odor memory and remembered the reward value of previously learned odor pairs after three weeks and one year of recess. Compared to other species tested previously on the same sets of odorants, the Asian elephants performed at least as good as mice and clearly better than human subjects, South African fur seals, squirrel monkeys, pigtail macaques, and honeybees. Taken together, these results support the notion that the sense of smell may play an important role in regulating the behavior of Asian elephants.

The sense of smell plays an important role in regulating the behavior of Asian elephants but until now, no behavioral test to systematically assess the olfactory capabilities of this species existed. Using a voluntary, food-rewarded two-alternative operant conditioning procedure, three female Asian elephants were successfully taught to discriminate between rewarded and unrewarded odors and also succeeded in intramodal stimulus transfer tasks in which either the rewarded odor, or the unrewarded odor, or both odors were exchanged simultaneously for new odors. The animals readily mastered the initial task within only 120 stimulus contacts, demonstrating rapid olfactory learning and performing at least as good as rodents and dogs and even better than other species, including nonhuman primates, tested in similar studies before. When presented with pairs of structurally related odorants, the discrimination performance of the elephants decreased with increasing structural similarity of the odorants, but the animals still significantly discriminated between aliphatic acetic esters even when they only differed by one carbon chain length. The elephants also demonstrated an excellent long-term odor memory and successfully remembered the reward value of previously learned odor stimuli after two, four, eight and even 16 weeks of recess in testing. The paradigm developed and applied in the present study proved to be useful to assess the olfactory capabilities in Asian elephants.

Captive populations of Asian elephants (Elephas maximus) in North America are not self-sustaining, and increasing reproductive success within captive populations is a high priority. The ability to accurately predict parturition can have a direct impact on elephant welfare. Elephants in captivity often require significant preparation and management throughout the birthing process, and complications during labor and delivery can necessitate immediate intervention, including stillbirth, protracted labor, maternal aggression towards a newborn calf, and dystocia. Being able to predict when parturition will commence can ensure appropriate staff is available and adequate monitoring is performed. Routine endocrine sampling can be used to predict parturition in Asian elephants, with a drop in progesterone (P4) to baseline levels signaling parturition in 2-5 days. However, we determined this method is not without limitations, and it is not used in all institutions that house elephants. As changes in hormones regulate and alter behaviors, we investigated behavioral indicators as an additional management tool for predicting parturition, a time of drastic hormone changes. We conducted a study of five pregnancies in Asian elephants at the Oregon Zoo, U.S.A, and Taronga Zoo, Australia, between 2008 and 2012. In Chapter 2, I evaluated progesterone (P4) and cortisol levels across three time periods: Baseline; Pre, (the week preceding the drop in P4); and Post, (the period after the P4 drop). Levels of P4 were significantly lower, and levels of cortisol were significantly higher in the days just prior to parturition. I found considerable intra- and inter-individual variation in both endocrine profiles, which can make endocrine assessments difficult to interpret in real time. In Chapter 3, I investigated whether behaviors in the preparturition period could be predictive of impending parturition in the Asian elephant. ANOVA results indicated a significant difference in the amount of time that elephants spent walking backwards across three time periods (F(2) = 3.723, p = 0.033), with the behavior increasing as parturition approached. These results were supported by a non-parametric Kruskal- Wallis. Using a generalized linear mixed model (GLMM), I found that as P4 levels decrease, walking backwards behavior significantly increases. In Chapter 4, I evaluated investigative trunk behaviors, or "trunk checks", directed towards the temporal gland near the ear, mammary glands, vulva and anus of the pregnant dam. Investigative behaviors included both self-directed behaviors and those sent from herd mates towards the pregnant dam. Self-directed behaviors are most likely associated with physical changes in the pregnant dam, such as using the trunk to pull on swollen teats. Other-directed behaviors may stem from chemo-sensory signaling or other types of communication between herd mates, such as detecting changes in progesterone or cortisol. I ran GLMM and found that four trunk-check behaviors varied significantly with P4 and/or cortisol profiles. These were: self-checks of mammary glands increased with decreasing P4 levels; herd-mate-checks of mammary glands increased with decreasing P4 levels; self-checks of vulva increased with decreasing P4 levels and increasing cortisol levels; herd-mate-checks of anus increased with increasing cortisol levels. In Chapter 5, I evaluated activity budget behaviors in the pregnant elephants. Generalized comparisons were made to published activity budgets of typical captive Asian elephants. I report that activity budgets are within the range of normal activity, though I note a high level of inter-individual variation. In addition, I compared two sampling techniques, including one-zero and instantaneous sampling, that were used for activity budget data collection. I discuss the different results obtained by each sampling technique. These results are a very promising indication that behaviors, including walking backwards and multiple trunk-check behaviors, are changing over time or with parturition-related hormone profiles. We recommend that keepers, veterinary staff, and other observers that are familiar with the regular behavioral repertoire of a pregnant female should pay close attention to these highlighted behaviors. Keeping track of these behaviors, especially in conjunction with P4 and cortisol tracking, can help staff refine existing windows of expected parturition.

African savanna elephants (Loxodonta africana) and Asian elephants (Elephas maximus) are a focus of welfare research in zoos due to their high intelligence, complex social structure, and sheer size. Due to these challenges, some argue that zoos are inherently incapable of providing appropriate care for elephants, while others believe that zoos can fulfill the needs of these species with improved husbandry. There is a general consensus from both within and outside of zoos, however, that zoos must improve their elephant programs or cease exhibiting these animals altogether. Now more than ever, applied research on zoo elephant welfare is needed to provide context for this debate. Researchers are interested in how far zoo elephants walk due to the potential health and welfare benefits of walking in these highly mobile species. Zoo researchers recently adopted GPS technology to study elephant walking, and preliminary evidence suggests that African elephants in large zoo exhibits walk distances that correspond with wild elephants under non-extreme conditions. However, data are limited from Asian elephants and from elephants in more typically-sized exhibits. In Chapter Two, I discuss important methodological considerations of utilizing GPS in a zoo environment, including an introduction to the technology, sources of error and mitigation, methods to improve GPS performance, and possible effects of GPS device attachment on animal behavior. This review shows GPS performance is adequate for tracking zoo elephant walking when proper methodological techniques are applied, and should serve as a useful reference for zoo researchers considering using GPS. In Chapter Three, I used GPS anklets to measure outdoor daily walking distance in 56 adult female African (n = 33) and Asian (n = 23) elephants housed in 30 zoos. I collected 259 days of data and found that elephants walked an average of 5.34 km/day with no significant difference between species. Multivariate regression models predicted that elephants with more dynamic feeding regimens (more diverse feeding types and frequencies; unscheduled feeding times) will walk more. Distance walked was also predicted to be higher in elephants that spend time in a greater number of different social groups. Distance walked was predicted to decline with age. Finally, I found a significant negative correlation between distance walked and nighttime space experience. The results of the analysis suggest that zoos that want to increase walking in their elephants need not rely solely on larger exhibits, but can increase walking by adding quality and complexity to exhibits. However, my results failed to establish a definitive link between walking distance and other validated measures of elephant welfare. Thus, the direct health and welfare benefits of walking in zoo elephants remain unresolved. Resting behaviors are an essential component of animal welfare, but have received little attention in zoological research. In Chapter Four, I used accelerometers in anklets to complete the first large-scale multi-species investigation of zoo elephant recumbence. I collected 344 days of data from 72 adult female African (n = 44) and Asian (n = 28) elephants at 40 zoos. I found that African elephants are recumbent an average of 2.14 hours/day, which is significantly less than Asian elephants at 3.22 hours/day. Multivariate regression models predicted that African elephant recumbence increases when they experience more space at night, and Asian elephant recumbence increases when they spend time housed alone. Both species showed a similar response to substrate, such that African elephants spending time on all-hard substrates are predicted to be recumbent less, while Asian elephants spending time on all-soft substrates are predicted to be recumbent more. The discovery that occasional non-recumbence is a common behavior in zoo elephants also introduces a new area of research that may have important animal welfare consequences. Finally, this study established that zoos should continue their efforts to replace hard substrate with soft substrate in order to provide zoo elephants with environments that facilitate recumbence. Overall, this work assessed walking and recumbence in zoo elephants, which will allow zoos to gauge the prevalence of these behaviors in their elephants as compared to the sub-population studied here. A variety of factors that are associated with these behaviors were also identified. With this information, zoos can prioritize modifications to their facilities and animal management programs to create an environment that encourages zoo elephants to express walking and recumbence behavior, should they choose to do so. This work is one component of the Elephant Welfare Project, the largest zoo animal welfare project ever undertaken, and is unprecedented in both scope and scale. The project was funded by the Institute of Museum and Library Services (IMLS), an independent, U.S., federal, grant-making agency that supports libraries, museums, and zoos. At the time of this writing, the first manuscripts from this project are being submitted to academic journals. These papers will describe the prevalence and distribution of a variety of elephant behaviors and welfare indicators, serve as a benchmark for future elephant welfare studies, and aid in decision making with regard to best practices in elephant management.

Stereotypic behaviors in all animals are more often than not associated with poor welfare. Limited access to perform species specific behaviors is often a reason for the development of stereotypies. Elephants with their great intelligence and need of social contact, coupled with a destructive a behavior are especially difficult to house in captivity. To decrease the occurrence of stereotypic behaviors in elephants, environmental enrichment in form of food enrichment is a good option, since elephants have a great need of foraging. The aim of this study was to investigate if an extra supply of food enrichment could decrease the presence of stereotypic behaviors at night in three Asian elephants at Kolmården zoo. Already existing hay nets attached to wires in the roof were used and connected to a timer. The hay nets were lowered to vision trunk reach between 6:00 am and 6:30 am during five days and the frequencies of selected behaviors were compared with the frequencies of the same behaviors during five preceding baseline nights. The animals were filmed using mounted cameras with IR lights. There was a significant decrease of stereotypic behavior for one of the elephants, but all three showed an increase in foraging whereof the increases were significant for two of them. One of the elephants showed no stereotypic behaviors at all during the study. To keep in mind is that the elephants have different backgrounds and experiences which might have influenced the results.

Stereotypes in captive elephants is a widespread issue. Ways to tackle this is with social and feedenrichment among others. The aim of this study was to investigate if extra feed supply wouldaffect stereotypic behaviour in Asian elephants held in captivity. Three elephant cows wereprovided with extra feed and recorded during the night and morning. The results showed asignificant decrease in stereotypic behavior for one of the elephants, from 31% to 9,5% (P =0,003). While the second elephant did not engage in stereotypic behaviour, the third elephantshowed the same frequency of stereotypic behavior, 9%, during both baseline and treatment. Thethird elephant did increase her foraging significantly, from 31% to 54% (P = 0,02). Waken timespent foraging increased for all three elephants to 64-80% which is in the same range as in wildAsian elephants. Additional enrichment in the form of extra feed supply can be used both todecrease stereotypic behaviours and to increase foraging.

This thesis presents an examination of the political ecology of human-elephant relationships in India. Its overall aim is to revitalize the ecology that has been sifted out from the discipline. The thesis draws upon, and consequently develops, more-than-human geography through a sustained engagement of nature-society relations in a non-Western context. The thesis has three broad objectives. First, to examine what more-than-human geography’s emphasis on non-dualistic forms of agency, could contribute to understandings of policy, planning and politics in conservation. Second, to examine the spatial dimensions of human-elephant relations and the social orderings of space which influence these relationships. The third objective of the thesis is to interrogate the politics of elephant conservation through a sustained engagement with diverse modes of human-elephant encounters and the socio-political assemblages with which they are entangled. The thesis first deploys and develops the concept of ‘encounter value’ to account for the different forms of human-elephant encounters and how they contribute to the political economies of biodiversity conservation. The thesis then draws from a multi-sited ethnography examining both encounters and spaces of elephant conservation. It shows how elephants help forge connections across difference and the ways their geographies are reconfigured by global networks of conservation. The third empirical section has an implicit spatial dimension. It is concerned with writing a ‘more-than-human’ geography of landscapes, examining how humans and elephants cohabit with and against the grain of political design. Finally, the thesis examines politics as an ecology of relations, showing how human-elephant relations as well as social and political outcomes may be mediated by materials. Modes of enquiry between these papers overlap. They offer critical insights into three themes that interface between political ecology and more-than-human geography. First, the thesis contributes to conceptualizing modes of human-animal encounters in a symmetrical fashion. It explicates the role of nonhuman agency as an organizing force in political economies of conservation. Second, it posits new understandings of the spaces of animals. This is developed in two ways: landscapes as dwelt, political achievements and as fluid spaces emerging through international networks of environmental governance. Third, the thesis ecologizes politics and goes beyond the humanist frameworks of political ecology. It fosters novel conversations between more-than-human geography and the postcolonial critique of political ecology in the context of human-elephant relationships. Taken together, the thesis offers up a concerted, symmetrical and novel approach to the study people’s relations with animals.

Animals in zoos live in more barren environments than their conspecifics in nature. In nature, elephants spend more than 75 % of the day foraging and have been observed lying down two hours during night. Stereotypies are common in many different species in zoos. Stereotypic behaviors may constitute up to 50 % of the daily activity budget of zoo elephants. Modern zoos try to reduce stereotypic behaviors by means of environmental enrichment. Three elephant cows at Kolmårdens Djurpark were observed during night to map their nocturnal behaviors and see how they changed with feeding enrichment. The elephants were lying down on their sides 25 % of the time observed. With feeding enrichment offered in the morning, the stereotypies was reduced in one individual from 43 % to 13 % (P<0.01) and the foraging was increased in two individuals from 45 % to 80 % (P<0.01) and from 45 % to 64 % (P<0.05). Increased food access increased foraging behavior and as environmental enrichment was able to reduce the stereotypic behaviors in these Asian elephant.

Environmental enrichment (EE) is used to improve the life of captive animals by giving them more opportunities to express species-specific behaviours. Zoo elephants are one of the species that is in great need of EE because their environment is often barren. Before making EE permanent, however, it is wise to test first if it works as intended, to save time and money. Maximum price paid is one measure that can be used to assess if an animal has any interest in a resource at all. Food is often used as a comparator against EEs in these kinds of studies. The aim was to investigate if the maximum price paid concept could be used to measure the value of EEs for the two female Asian elephants at Kolmården and to find an operant test suitable for them for the experimental trials. Three series of food trials were done with each elephant, where they had to lift weights by pulling a rope with their mouth to get access to 5kg hay. The elephants paid a maximum price of 372 and 227kg, respectively. However, the maximum price the elephants paid for access to the hay was not stable across the three series of trials. Hence it is recommended that the comparator trials are repeated close in time to the EEs to be tested. The readiness by which these elephants performed the task makes it worthwhile to further pursue this approach as one of the means to improve the well-being of zoo elephants.

Translocation of wild Asian elephants (Elephas maximus) is used extensively to mitigate human-elephant conflict (HEC) in Peninsular Malaysia since 1974. Very little is known about the fate of translocated elephants after relocation due to challenges in observing elephants in the dense rainforest. Advances in wildlife endocrinology suggest that faecal glucocorticoid metabolites (fGCM) can be used to study adrenal activity remotely, to assess the Hypothalamic-Pituitary-Adrenal (HPA) axis response towards stressors. The aim is to assess the impact of translocation on wild Asian elephants in Peninsular Malaysia using faecal endocrinology and GPS technology. The specific objectives are: (i) adapting hormone sampling methods for use under tropical field conditions, (ii) comparing fGCM concentrations between translocated and local resident elephants using enzyme immunoassay, and (iii) quantifying gastrointestinal parasite eggs and microflora ciliates in faecal samples to detect signs of immunosuppression. We found that Asian elephant’s fGCM (80 dungpiles, 685 subsamples) are stable up to eight hours in the field. From the monitoring of wild elephants at the release sites, between two months up to a year, translocated elephants (N=5) had lower fGCM concentrations in comparison to local resident elephants (N=4; Linear Mixed Models: t=-2.77, df=7.09, P=0.027). There were no differences in gastrointestinal parasite egg counts (P > 0.05) or microflora ciliate counts (P > 0.05) between translocated and local resident elephants. In conclusion, translocation does affect elephant physiology but this is in the opposite direction from that expected – a prolonged decrease rather than increase of adrenal activity. It is unknown if these conditions could cause immunosuppression, but it could adversely affect stress response and health of the elephant (e.g. adrenal insufficiency, chronic fatigue or Post-Traumatic Stress Disorder). When assessing HEC mitigation, conservation authorities and other stakeholders need to consider that translocation may not be the best solution for HEC, as it will have long-term consequences on elephants’ health.

Asian elephants utilize two chemical signals that have been described to function in reproduction: (1) (Z)-7-dodecenyl acetate (Z7-12:Ac) is released by females near ovulation, and (2) frontalin is released by males around the time of musth. Signaling theory posits that the concentration at which either compound is emitted should have implications for the response of the receiver, varying with factors such as sex and reproductive experience. Here, the objectives were to: (1) investigate the effect of concentration on receiver chemosensory behavior in an effort to identify detection thresholds and concentrations of maximum response for reproductively experienced or inexperienced male and female Asian elephants, and (2) characterize the broader behavioral impacts of each of these compounds in an effort for application as environmental enrichment in captive settings. Concentrations from 0.0 mM to 2.0 mM of both frontalin and Z7-12:Ac were bioassayed simultaneously with captive elephants housed at facilities across North America in two experiments: one that tested mid-range concentrations and a second that tested low and high concentrations. There was a general increase in chemosensory response with increasing concentration of both compounds regardless of sex or reproductive experience. Females exhibited a lower detection threshold for frontalin, and the opposite was true for males with Z7-12:Ac. Reproductive experience also influenced thresholds: inexperienced males had a higher threshold than experienced males for frontalin (the same was true for females), and experienced males were able to detect Z7-12:Ac samples as low as 10–7 mM. Aside from inexperienced males, all elephants responded maximally to the 1.0 mM samples of both compounds. Elephants exposed to mid-range concentrations of either compound showed no notable changes in behavior after application of the signals, although inexperienced males spent less time inactive and more time walking after frontalin bioassays, and inexperienced females foraged more after exposure to Z7-12:Ac. Interpreted together, this suggests that the concentration at which either compound is emitted has strong implications for chemosensory response based on the identity of the receiver in Asian elephants, although it is unclear whether these compounds have other behavioral effects that can be targeted for a goal-oriented olfactory enrichment program.

付記する学位プログラム名: 霊長類学・ワイルドライフサイエンス・リーディング大学院
Kyoto University (京都大学)
0048
新制・課程博士
博士(理学)
甲第22298号
理博第4612号
新制||理||1661(附属図書館)
京都大学大学院理学研究科生物科学専攻
(主査)教授 幸島 司郎, 教授 平田 聡, 教授 伊谷 原一
学位規則第4条第1項該当

International organisations such as the World Health Organisation highlight the road crash problem in less motorised (or developing, or low income) countries like those in Southeast Asia and recommend the adoption of Western road safety measures. However, there are many differences between highly motorised and less motorised countries which raise questions about how successfully Western road safety knowledge and expertise can be transferred.----- A review of the statistical information on road crashes shows a great deal of uncertainty about both the scale and likely trajectory of road fatalities globally, in less motorised countries and in Asia. It is generally agreed, however, that Asia accounts for around half of all road fatalities, and analysis of the limited available data shows both that Southeast Asia is not an atypical region of Asia in road safety terms, and that Thailand and Vietnam are not atypical of Southeast Asian countries.----- A literature review of recommended practice approaches to road safety transfer in Asia shows that there are many economic, institutional, social and cultural factors which potentially influence the success of transfer. The review also shows that there is no coherent, comprehensive approach which either conceptualises these factors and their relationship to transfer outcomes, or uses an analysis of these factors to plan or modify transfer. To address this gap, this thesis develops a 'road safety space' model as a tool for conceptualisation and analysis, based on a biological metaphor which views the transfer of road safety measures from one context to another as analogous to the transfer of a species into a new ecological space. The road safety space model explicitly considers economic, institutional, social and cultural factors (from specific to broad) which influence the particular road safety issue which a particular road safety transfer effort seeks to address. A central contention of this thesis is that the road safety space model is both a feasible and useful tool to improve the process of road safety transfer to less motorised countries. Road safety space analysis is seen to have a role in a broader process of selection of road safety measures for transfer, along with knowledge of how the measures are considered to operate.----- The research reported in this thesis is comprised of three studies. Study 1 reviewed evaluations of road safety transfer to Thailand and Vietnam. Studies 2 and 3 were case studies of road safety transfer to Thailand and Vietnam respectively.----- Study 1 was an analysis of existing evaluations of road safety transfer to Thailand and Vietnam. The aims were to analyse the evaluations for their consideration of contextual factors, as described in the road safety space model, and to discuss whether the road safety space model assisted in understanding the reasons for success or failure of transfer. However, very few such evaluations exist, and those that were found generally lacked information on whether contextual factors were considered. This indicated the need for a more detailed, in-depth qualitative investigation of particular cases of road safety transfer, in order to investigate the feasibility and utility of the road safety space model.----- Two case studies (Study 2 and Study 3) were conducted to test whether the road safety space approach was both feasible and useful as a means of improving road safety transfer efforts. Study 2 was a case study of the development and implementation in Thailand of a road safety education program for school children, which involved the transfer of Western research and techniques. The transfer agents (i.e. those who effected the road safety transfer) were Australian consultants working for the Australian Road Research Board (ARRB). The transfer was funded by the World Bank and managed by the Thai Ministry of Education (MOE). Study 3 was a case study of the development and implementation of a motorcycle helmet wearing program in Vietnam, which involved the transfer of Western knowledge, techniques and technology. The transfer agents were staff of Asia Injury (AI), a non-government organisation (NGO), and the program was funded initially by a charitable fund, with the intention of becoming self-funding through operation of a helmet factory.----- The case studies employed background research into existing information on economic, institutional, social and cultural factors relevant to the road safety issues (road use behaviour of school children in Thailand and motorcycle helmet purchase and wearing in Vietnam), and collected data through interviews with key informants, analysis of secondary sources and observations. This information was used to derive the road safety space for each road safety issue, to identify the road safety space recognised and addressed by the transfer agents (ARRB and AI), and to determine which factors they missed, or were aware of but took no action on. The focus of this analysis was on the processes used in transfer, not on the road safety outcomes of transfer, although these provided information on the processes as well. Available evaluation information was used to draw links between the omissions and the success of the transfer processes. It was noted that information on how the transferred measures operate should come from a road safety space analysis in the originating country, although this raised questions about selection of country and time (when the measure was first introduced, or in its maturity).----- The feasibility and utility of the road safety space model were discussed. It was clear that the model provided information on the cases which was missed by the transfer agents. The questions examined next were whether this information could have been obtained from an exercise conducted before the transfer had commenced, whether the required effort and cost justified the potential benefits, and whether the information on the road safety space could have been useful for the transfer agents. Comparisons between the road safety spaces for the two cases showed some areas of commonality, e.g. perceptions of police corruption, but also many differences. It was considered likely that some broad factors could be generic, and the possibility was mooted that less motorised countries share issues with police enforcement. This requires further research, however, and at this stage it is better to treat each road safety space as a unique combination of contextual factors influencing the road safety issue of interest.----- It is concluded that the road safety space model is feasible if used in such a way as to minimise the research involved, and useful, although the degree of utility needs to be further explored in a prospective study. The limitation introduced by restricting informants to those who could speak English are discussed. An approach using road safety space analysis is recommended, emphasising analysis of the country to which the road safety measure is being transferred, supplemented by analysis of the originating country road safety space. Gaps in knowledge are identified for further research and development, in particular the theoretical and practical understanding of road use behaviours and their modification in less motorised countries in Southeast Asia. Elaboration of the model is also recommended, to take into account the influence of the type of measure transferred, the role of the transfer agent, the area of road safety (education, engineering or enforcement), and the time dimension (the time which might be needed for a transfer to show its effects).----- The findings of this research are likely to be applicable to road safety transfer in other less motorised regions of the world, however prospective testing is needed. They may also be relevant to issues of transfer for areas other than road safety, in particular public health and traffic engineering, where similar economic, institutional, social and cultural issues come together.

International organisations such as the World Health Organisation highlight the road crash problem in less motorised (or developing, or low income) countries like those in Southeast Asia and recommend the adoption of Western road safety measures. However, there are many differences between highly motorised and less motorised countries which raise questions about how successfully Western road safety knowledge and expertise can be transferred.----- A review of the statistical information on road crashes shows a great deal of uncertainty about both the scale and likely trajectory of road fatalities globally, in less motorised countries and in Asia. It is generally agreed, however, that Asia accounts for around half of all road fatalities, and analysis of the limited available data shows both that Southeast Asia is not an atypical region of Asia in road safety terms, and that Thailand and Vietnam are not atypical of Southeast Asian countries.----- A literature review of recommended practice approaches to road safety transfer in Asia shows that there are many economic, institutional, social and cultural factors which potentially influence the success of transfer. The review also shows that there is no coherent, comprehensive approach which either conceptualises these factors and their relationship to transfer outcomes, or uses an analysis of these factors to plan or modify transfer. To address this gap, this thesis develops a 'road safety space' model as a tool for conceptualisation and analysis, based on a biological metaphor which views the transfer of road safety measures from one context to another as analogous to the transfer of a species into a new ecological space. The road safety space model explicitly considers economic, institutional, social and cultural factors (from specific to broad) which influence the particular road safety issue which a particular road safety transfer effort seeks to address. A central contention of this thesis is that the road safety space model is both a feasible and useful tool to improve the process of road safety transfer to less motorised countries. Road safety space analysis is seen to have a role in a broader process of selection of road safety measures for transfer, along with knowledge of how the measures are considered to operate.----- The research reported in this thesis is comprised of three studies. Study 1 reviewed evaluations of road safety transfer to Thailand and Vietnam. Studies 2 and 3 were case studies of road safety transfer to Thailand and Vietnam respectively.----- Study 1 was an analysis of existing evaluations of road safety transfer to Thailand and Vietnam. The aims were to analyse the evaluations for their consideration of contextual factors, as described in the road safety space model, and to discuss whether the road safety space model assisted in understanding the reasons for success or failure of transfer. However, very few such evaluations exist, and those that were found generally lacked information on whether contextual factors were considered. This indicated the need for a more detailed, in-depth qualitative investigation of particular cases of road safety transfer, in order to investigate the feasibility and utility of the road safety space model.----- Two case studies (Study 2 and Study 3) were conducted to test whether the road safety space approach was both feasible and useful as a means of improving road safety transfer efforts. Study 2 was a case study of the development and implementation in Thailand of a road safety education program for school children, which involved the transfer of Western research and techniques. The transfer agents (i.e. those who effected the road safety transfer) were Australian consultants working for the Australian Road Research Board (ARRB). The transfer was funded by the World Bank and managed by the Thai Ministry of Education (MOE). Study 3 was a case study of the development and implementation of a motorcycle helmet wearing program in Vietnam, which involved the transfer of Western knowledge, techniques and technology. The transfer agents were staff of Asia Injury (AI), a non-government organisation (NGO), and the program was funded initially by a charitable fund, with the intention of becoming self-funding through operation of a helmet factory.----- The case studies employed background research into existing information on economic, institutional, social and cultural factors relevant to the road safety issues (road use behaviour of school children in Thailand and motorcycle helmet purchase and wearing in Vietnam), and collected data through interviews with key informants, analysis of secondary sources and observations. This information was used to derive the road safety space for each road safety issue, to identify the road safety space recognised and addressed by the transfer agents (ARRB and AI), and to determine which factors they missed, or were aware of but took no action on. The focus of this analysis was on the processes used in transfer, not on the road safety outcomes of transfer, although these provided information on the processes as well. Available evaluation information was used to draw links between the omissions and the success of the transfer processes. It was noted that information on how the transferred measures operate should come from a road safety space analysis in the originating country, although this raised questions about selection of country and time (when the measure was first introduced, or in its maturity).----- The feasibility and utility of the road safety space model were discussed. It was clear that the model provided information on the cases which was missed by the transfer agents. The questions examined next were whether this information could have been obtained from an exercise conducted before the transfer had commenced, whether the required effort and cost justified the potential benefits, and whether the information on the road safety space could have been useful for the transfer agents. Comparisons between the road safety spaces for the two cases showed some areas of commonality, e.g. perceptions of police corruption, but also many differences. It was considered likely that some broad factors could be generic, and the possibility was mooted that less motorised countries share issues with police enforcement. This requires further research, however, and at this stage it is better to treat each road safety space as a unique combination of contextual factors influencing the road safety issue of interest.----- It is concluded that the road safety space model is feasible if used in such a way as to minimise the research involved, and useful, although the degree of utility needs to be further explored in a prospective study. The limitation introduced by restricting informants to those who could speak English are discussed. An approach using road safety space analysis is recommended, emphasising analysis of the country to which the road safety measure is being transferred, supplemented by analysis of the originating country road safety space. Gaps in knowledge are identified for further research and development, in particular the theoretical and practical understanding of road use behaviours and their modification in less motorised countries in Southeast Asia. Elaboration of the model is also recommended, to take into account the influence of the type of measure transferred, the role of the transfer agent, the area of road safety (education, engineering or enforcement), and the time dimension (the time which might be needed for a transfer to show its effects).----- The findings of this research are likely to be applicable to road safety transfer in other less motorised regions of the world, however prospective testing is needed. They may also be relevant to issues of transfer for areas other than road safety, in particular public health and traffic engineering, where similar economic, institutional, social and cultural issues come together.

Darwin was troubled by elaborate male traits observed in many species that are seemingly maladaptive for survival, the peacock’s tail being the most iconic of all. He wrote "The sight of a feather in a peacock’s tail, whenever I gaze at it, makes me sick" because it challenged his theory of evolution by natural selection for adaptive traits. The extreme length of the tail may render a peacock more vulnerable to predation and therefore maladaptive for survival. To account for the evolution of apparently maladaptive traits he proposed the theory of sexual selection, wherein, traits that directly enhance mating success may be selected for, either as weapons in male-male competition for mates or as ornaments preferred by females. Male and female elephants in the proboscidean evolutionary radiation have had tusks and show extreme exaggeration in size and form. However, tusk in the Asian elephant (Elephas maximus) is sexually dimorphic as it is expressed only in the males, hinting at a possibility that opposing selection (sexual selection advantage to males and natural selection disadvantage to females) may have been the processes behind this pattern of tusk expression. Intriguingly, tuskless males (male dimorphism with respect to tusk) also occur at fairly high frequencies in some Asian elephant populations (∼50% in norteastern India and ∼95% in Sri Lanka). Theory states that dimorphic males can also occur in a population in stable frequencies as a consequence of sexual selection. I explored sexual selection on elephant tusks as possible mechanism leading to the observed patterns of tusk dimorphism in the elephants. All elephant populations on earth have been harvested for ivory, therefore, artificial selection (selective poaching of tusked elephants for ivory) is another possible cause of tusk dimorphism. I developed mathematical models of population genetics, population dynamics and conducted field observations of mating behavior of Asian elephant in Kaziranga National Park, Assam to understand the evolution of tusk dimorphism in elephants. Darwin’s sexual selection theory was controversial when proposed in 1871 and continues to remain so in 2014. In the introduction of my thesis I have discussed Darwin’s two classical mechanisms of sexual selection, namely, male-male combats for mates and female mate choice based on male traits. The latter was viewed with considerable skepticism by his con-temporary Alfred Russell Wallace and more recently deemed "fundamentally flawed" by Joan Roughgarden. Therefore, I have also discussed the arguments against female mate choice for male traits found in literature. I have reviewed current knowledge about sexual selection for sexually dimorphic male traits of body size and musth, in the African and Asian elephant and state why I have hypothesized that tusks may also be under sexual selection. Sexually selected traits are expected to be genetically determined, therefore, I explored mathematically (Chapter 1) the genetic basis of evolution of sexual dimorphism. Fisher proposed that sexually selected male display traits originate in both the sexes but are suppressed in the females by modifier genes, when the trait becomes deleterious to females. Thus, sexually antagonistic selection on a trait and sex-specific gene expression can lead to the evolution of sexual dimorphism. Tusk is sexually monomorphic in the probocideans that are ancestral to both the African (Loxodonta africana) and Asian elephant (Elephas maximus). Tusk continues to remain monomorphic in the African elephant but has become sexually dimorphic in the Asian elephant. Tusk, therefore could be a sexually selected male trait that evolved according to the Fisherian model. Intriguingly, tuskless males occur at very high frequencies in some Asian elephant populations. The tusked and tuskless male morphs could be alternate male mating strategies, occurring at evolutionarily stable frequencies. Alternatively, the observed male tusk dimorphism, could be a consequence of artificial selection against tusked individuals, due to selective harvest of tusked males. Furthermore, male African elephants are more intensely poached for ivory than female elephants. Yet the frequency of tuskless individuals has increased more rapidly among females than in males. In essence, sexual dimorphism could be evolving among such poached populations. Is such rapid, contemporary evolution of sexual dimorphism, possible through the Fisherian modifier gene mechanism? A 2-loci genetic model (with X-linked trait gene and an autosomal modifier gene) (Rice 1984), a slight variant of the model (with X-linked modifier gene, and an autosomal trait gene) and an entirely autosomal model, were analyzed for the rate of evolution of sexual dimorphism, under different selection pressures for tusk possession. Negative frequency dependent selection was introduced into the model of tusk evolution in accordance with Gadgil’s model for the evolution of male dimorphism as consequence of sexual selection (Gadgil 1972). In two of the 2-loci models (in which tusk gene in autosomal), tusklessness evolved much more rapidly in females than in males, under equal negative selection pressures. The models predict several combinations of time-lines and negative selection pressures for effecting a particular change in the frequency of tusklessness. Model predictions were com-pared with observed changes in the frequency of female tusklessness, in one South Ugandan, African elephant population (∼2% to 10% in 5 to 9 generations) and male tusklessness (∼5% to 50% in 25 to 40 generations) in one north eastern Indian, Asian elephant population. The models predict strong selection pressures of 30% to 50% reduction in fitness, that can effect an 8% increase in tusklessness, in the African elephant population, within time-lines of 9 to 5 generations (∼225 to 125 years) respectively. For the male Asian elephants, natural selection against tusked males on an already sexually dimorphic population, must have been in operation and shifted the population to 5% male tusklessness. The models predict that artificial selection with 20 to 30% fitness cost to tusked males, operating for 40 to 25 generations (∼1000 to 600 years) respectively, can further shift the population from ∼5% to ∼50% tusklessness. Asian elephant populations may already have been in a transient phase of evolution, tending towards tusklessness, with recent artificial selection hastening the process. The two major pre-dictions from this modeling exercise are (1) artificial selection could have played a significant role in the evolution of male tusk dimorphism in the Asian elephant (2) a lack of or very mild current sexual selection on tusks in the male Asian elephant. Both these predictions may be empirically verified. Chapters 2 and 3 are attempts at empirical verification of prediction (1) and Chapters 4 and 5 of prediction (2). From historical references to elephant harvest in Assam, we do know that artificial selection has been in operation, but whether it has played a major role in causing male tusk dimorphism needs to be established. It may be possible to detect signatures of significant past harvest from current demographic structure of an elephant population. Sustained biased harvesting of a particular sex and or age class from an animal population alters the sex ratio and age structure (relative proportion of individuals in each age and sex class) of a population considerably (Sukumar 1989). It may be possible to back infer the harvest scenario by studying the deviation of current age and sex ratios from natural age and sex ratios. In Chapter 2, I explored models of population dynamics under different harvest regimes and its effect on age and sex ratios. I described a method to infer unknown harvest rates and numbers from age and sex ratios, 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 deter-mined from the history of harvest and an estimate of population size. I validated this model with published data on age and sex ratios of one Asian and African elephant population with fairly reliable data on elephant harvest as well. In Chapter 3, I applied this model to the demographic data that I collected from a wild Asian elephant population in Kaziranga National Park, Assam, India (where more than 50% of the adult males are tuskless). Male polymorphism of sexually-selected male traits occur at stable frequencies in populations of several species. The different male morphs of the trait are hypothesized to be alternate male mating strategies with equal life time reproductive fitness. Male Asian elephants of Kaziranga National Park, Assam are dimorphic with respect to tusk possession: ∼50% of the males are tuskless (and are locally called makhnas). Makhnas could be trading tusk for either longevity, larger body size, testicular volume and or duration of musth as alternate mating strategies. On the other hand makhnas may have increased to a very high frequency primarily due to selective removal (captures for domestication and hunting for ivory) of tusked males from the population for centuries. The aim of Chapter 3 was to examine the role of artificial selection in the evolution of makhnas. Prolonged male-biased harvest(removal from the population) is bound to alter the demographic structure of the population and leave a signature of the intensity and type of harvest on the residual population structure. The Kaziranga elephant population was considered as representative of elephant populations of north east India; A harvest modeling approach (described and validated in Chapter 2) was used to infer unknown harvest of elephants from demographic parameters estimated by sampling this elephant population during 458 field days in the dry season months of 2008–2011. The Kaziranga elephant population appears to have been harvested approximately for the past 700 to 1000 years with adult tusked males being harvested at approximately twice the rate of adult tuskless males, adult females and their immature offspring of both the sexes. The currently observed high frequency of tuskless males in Kaziranga therefore, may be a consequence of sustained artificial selection against tusked males for several centuries. The previous two Chapters have only examined some mechanisms for the loss of tusks in elephants. I proceeded to examine the possibility of evolution of tusks through Darwin’s mechanisms of male-male competition for mates and female mate choice. Elephant tusks are cited as an example of a male trait that has evolved as a weapon in male-male combats. In Chapter 4 I examined the role of 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. I 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 played 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. If not a weapon, tusks could be a male ornament that female elephants find attractive. I explored the interplay of the three male traits (body size, musth and tusk), male mating strategies and female mate choice in Chapter 5. In some species males obtain mating opportunities by harassment of females. Given the striking size difference between an adult male and female elephant, with males weighing at least 30% more than females, male coercion of females to mate is a possibility. A detailed study of the courtship behavior revealed that overt male harassment of females is rare and the ability of a male to mount and stay mounted on a female for copulation is under female control. Therefore female Asian elephants can exercise choice to mate but this is subtly different from exercising mate choice itself. Age-related male mating strategy (reported for the first time in the Asian elephant) exists in the Kaziranga elephant population and this strategy limits the ability of females to exercise choice. Young males (<25 years) predominantly show a sneak mating strategy. Middle-aged males (25–40 years), when in musth, mate–guarded oestrous females from sneakers and attempted mating but sometimes resorted to sneak mating when out of musth. Old males (> 40 years) attempted mating only during their musth phase and were seldom sneakers. Large/musth males received positive responses from estrous females towards courtship attempts significantly more often than did small/non–musth males. Tusked non–musth males attempted courtship significantly more often than did their tuskless peers, and had a higher probability of receiving positive responses than did tuskless males. A positive response, however, may not translate into mating because of mate–guarding by the dominant male. Females permitted large/musth males to stay mounted significantly longer than small/non-musth males. Musth and large body size may be signals of male fertility. Female mate choice in elephants thus seems primarily for traits that signal direct benefits of assurance of conception. Tusked males may attain sexual maturity faster than tuskless males. Therefore it is worth exploring if tusks function as signals of male fertility when males are young (15 to 25 years); this may be possible through hormonal and behavioral profiling of young tusked and tuskless males from 10 to 20 years of age. Overall all musth and body size appear to play a larger role in enhancing male mating success than tusks. Tusked males appear to have a weak sexual selection advantage (male-male domi-nance and female preference) over their tuskless peers, only in the young age class (15 to 25 years) in this population. Males in this age age class, seldom come into musth that would over-ride tusk as a signal of male dominance. Current sexual selection on tusks in this population, appeared to be insignificant and this may be verified through genetic analysis of paternity success. An important implication of musth and body size being stronger determinants of mating success than tusk possession is that, it could facilitate rapid evolution of tuskless males in the population under artificial selection against tusked individuals, even in a slow breeder such as the elephant. Musth may have evolved much later than tusks in elephants, therefore it is possible that tusks evolved under sexual selection before musth evolved. However, body size, in mammals in gen-eral appear to be under both natural and sexual selection. Gould has shown that the absurdly large and palmate antlers of the extinct Irish elk, scales allometrically with body size (Gould & Lewontin 1979). Phylogenetic studies of elephant evolutionary radiation indicate a general trends towards increase in body-size with size reduction and tendency towards dwarfism occurring only in island habitats (Palombo 2001). Tusk development, which is essentially tooth development may be closely linked to cranium development. Cranium development in turn may be linked to body size through allometric scaling laws. If so, any selection on body size is bound to act on tusk size. I propose that the evolution of elaborate tusks seen in elephants is primarily due to natural and or sexual selection acting on body size, and tusk just hitched a ride with body size. Tusks may be maintained in spite of tuskless males occurring in the population only because of a rather weak sexual selection advantage to tusk possession in contests in which males are symmetrical with respect to body size and musth status.

Masters Research - Master of Philosophy (MPhil)
Human-elephant conflict (HEC) in the form of crop-raiding, is a major conservation challenge to the long-term survival of elephant populations, simultaneously threatening the livelihoods and personal safety of people living in proximity to elephants. The widespread problem of HEC has led to a great deal of research into the causes, consequences and predictors of elephant crop-raiding activity. However, despite similarities across HEC situations, site-specific differences are also apparent. Furthermore, most studies focus on one facet of HEC when it is a complex issue requiring understanding of local elephant behaviour, identification of the characteristics and patterns of crop-raiding at the local scale, and careful implementation and monitoring of mitigation strategies. In this study, I selected a region of Sri Lanka experiencing high levels of HEC and sought to provide an in-depth assessment of the site-specific situation generated over a three-year period. Specifically, we aimed to: identify general patterns of behaviour occurring in local areas representing differing levels of anthropogenic disturbance to elephants; profile patterns and predictors of crop-raiding activity in a village heavily impacted by HEC; and test the effectiveness of beehive fencing as an Asian elephant deterrent tool. First, I provide initial evidence that elephants inhabiting areas of ‘medium’ level anthropogenic disturbance outside of protected boundaries, interrupt feeding and increase ‘reactive’ behaviours such as smelling and holding the ‘vigilance’ posture in response to immediate anthropogenic threats in the environment. In the absence of any known disturbances, there was no difference in general behaviours of male or female elephants between the two risk zones. Secondly, I show that elephant crop-raiding in Dewagiriya Village occurs year-round and follows no clear seasonal patterns. Similar to other HEC situations, male elephants are the predominant crop-raiders, and crop-raiding occurs almost exclusively at nights. Within-site variations in crop-raiding intensity were also identified, with properties closest to water tanks and forest habitat the most vulnerable. Finally, our three-year beehive fence trial showed that households using beehive fences around their gardens had significantly less elephant visits into their gardens then households without. Still, difficulties in attracting natural colonies, poor honey production, set-up costs, and farmer motivations were barriers to success. This study contributes to the general body of knowledge on elephant behaviour in an anthropogenically influenced contexts, and specifically on patterns of crop-raiding and mitigation efforts. Further research into the potential of beehive fences as an Asian elephant deterrent, preferably in a location more amenable to beekeeping, would help to determine the value of expanding this technique further in Sri Lanka, and elsewhere in Asia.

An animal’s ability to find thermal comfort can be can be limited within a zoo enclosure. This creates a need to design comfortable microclimates for the animal within their enclosure space. This is an exploratory study intended to investigate the relationships between an elephant’s movements throughout their space and the microclimatic conditions within its zoological enclosure. By outlining an understanding of microclimates and how they are affected by the landscape combined with knowledge of elephant behaviours it will be possible design or plan for microclimatic use within zoo enclosures. Observational data were collected from five Asian elephants (Elephas maximus). The movements of the elephants were monitored along with the use of microclimates within their enclosure. Thermal regulatory movements involving microclimatic use typically involved water and shade use, however evidence suggests that the animals were also avoiding terrestrial radiation from some of the gunite walls. It is important that animals are given the opportunity to find microclimates within the space to regulate their body temperature. Designing landscapes that incorporate microclimates within their habitats is crucial to the welfare and health of zoo elephants.

碩士
國立中興大學
獸醫學系暨研究所
107
Different hormone degradation responds differently to storage time and temperature in different species. The effects of storage time duration and temperature on hormone degradation before centrifugation have not been evaluated in elephant blood. We designed two project in this experiment. A total of 52 ml blood was collected from each elephant (Baanchang elephant camp, Chiang Mai), each project took up half of the blood. In Project EDTA, 26 ml of blood were divided into 13 aliquots and was left in EDTA tubes before centrifugation. 1 aliquot was centrifuged immediately and set as baseline (T0). 12 aliquots remained were divided into 3 temperature subgroups: group LT (4 ℃), group RT (25 ℃) and group HT (37 ℃), each group was incubated at relative temperature. One aliquot of each group was centrifuged and plasma was collected at 6hr, 24hr, 48hr, 62hr. In Project NA, 26 ml blood was divided into 13 aliquots and was left in non-anticoagulant tubes before centrifugation. All aliquots were treated like as Project EDTA. All the plasma and serum extracted was kept at -20 ℃ until analysis. Enzyme-linked immunoassay (EIA) was applied to measure the concentration of cortisol (male and female elephant), testosterone (male elephant) and progesterone (female elephant). Raw data was calculated as % of Time 0 (T0). One-way ANOVA and Dunnett’s test were used to analyze % of T0, to determine the effect of time, temperature and time * temperature. Significant difference was confirmed when P 〈0.05. The results were 1) cortisol, progesterone and testosterone in Project EDTA was stable at all temperature condition for up to 62 hours; 2) cortisol in Project NA decreased significantly after 48 and 62 hours in group HT (P 〈0.05 and P 〈0.01, respectively); 3) testosterone in Project NA decreased significantly after 48 and 62 hours in group HT (P 〈0.01 and P 〈0.001, respectively); 4) temperature and time duration before centrifugation did not affect concentration of progesterone in Project NA. These finding suggested that 1) plasma sample may provide better hormone stability at 37 ℃; 2) progesterone is the most stable hormone in elephant blood sample, and can be kept in tubes before centrifugation for up to 62 hours. 3) cortisol and testosterone concentration should be measured before 48 hours.

Sumatran elephant (Elephas maximus sumatranus) continue to decline due to habitat loss, poaching and conflict with humans. Yet, developing effective land conservation strategies for elephants is difficult because there is little information available on their foraging ecology, habitat use, movements and home range behaviors. Using the lead animal technique, 14 free-ranging, tame elephants at the Seblat Elephant Conservation Center (ECC) were observed for 4,496 hours to describe their foraging ecology and diet. The majority of their daily activity was feeding (82.2 ± 5.0%), followed by moving (9.5 ± 4.0 %), resting (6.6 ± 2.1%) and drinking (1.7 ± 0.6%), and individual activity budgets varied among individuals for all activities. At least 273 plant species belonging to 69 plant families were eaten by elephants and five plant families of Moraceae, Arecaceae, Fabaceae, Poaceae, and Euphorbiaceae were most commonly consumed. Elephants browsed more frequently than grazed, especially in the wet season. Levels of crude protein, calcium, phosphorus and gross energy in plants eaten by elephants in Seblat appeared adequate for meeting the nutritional requirements. Home range size of an adult female elephant in the SECC during 2007–2008, was 97.4 km2 for the MCP and 95.0 km2 for the 95% fixed kernel. There were no relationships between average monthly elephant home range sizes or movement distances with rainfall. Distances to rivers and ex-logging roads had little effect on elephant movements, but vegetation productivity, as measured by the Enhanced Vegetation Index, did affect elephant movements. We used resource selection and compositional analysis habitat ranking approaches to describe adult female elephant habitat use in the SECC. The elephant used medium canopy and open canopy forests more than expected; however, during the day closed canopy forests were used more than at night. Locating and capturing wild elephants in tropical rainforest environments are difficult and high-risk tasks. However, using tame elephants improves the search efficiency of finding wild elephants in dense forests and reduces risks to staff and target elephants. Use of experienced veterinarians and standing sedation techniques also greatly reduce the risks of elephant injury while immobilizing elephants.

Various ecological and human-induced disturbances play an important role in defining the health of an animal. To cope up with such threats or challenges to its homeostasis, an animal responds by secreting stress hormones (glucocorticoids) to mobilize the energy. In general, glucocorticoids help animals to cope up with the ‘stressors’. Prolonged exposure to any ‘stressors’ may lead to an elevation in the levels of glucocorticoids, thereby leading to reproductive inhibition, immune system suppression, neuronal death and impaired cognitive function and hence, threatening the survival and fitness of an animal. However, the long-term effects of environmental or anthropogenic disturbances on the fitness and survival are difficult to analyze in a slow reproducing, long-lived species such as elephants. Spread over 3% of India’s geographical area, India shelters around 28,000 to 30,000 Asian elephants (Elephas maximus; Bist, 2002; Santiapillai and Sukumar, 2006). Protected areas in India are estimated to form 22% of elephant habitat; while the rest of the elephant habitats extend outside the protected areas (Lenin and Sukumar, 2011) making them vulnerable to human-induced threats. Consequently, there are increased incidences of human-elephant conflicts leading to crop and property depredation and human-elephant mortality. Most of the studies, in context of such conflicts, highlight the temporal, spatial scales of conflicts majorly focusing on mitigation measures. None of the studies so far have elucidated how escalating human-elephant conflicts would influence the physiological health of Asian elephants. This study, hence, focused on addressing the proximate causation of the stress-response in free-ranging Asian elephants of the Bandipur National Park, the Nagarahole National Park and Hassan district of Karnataka, using a non-invasive technique for measuring faecal glucocorticoid metabolites (fGCM). There are several factors which could act as possible stressors to the physiological state of an elephant, but depending upon the feasibility and time limitations of conducting the study, we selected those factors which we thought are essential to address the proximate causation of stress-response in Asian elephants. Being the first detailed study to explore the stress-response in free-ranging Asian elephants, this study attempted to understand the influence of some of the fundamental ecological (seasonality, group size, and body condition) and human-induced stressors. The main body of the dissertation is divided into six chapters. The first and the last chapters discuss the general introduction and conclusion while the rest four chapters highlight the main four objectives of the study. The main objectives of the thesis were: 1. The first objective of this thesis was to validate and standardize some of the crucial parts of protocols to avoid any technique-based bias while interpreting the levels of faecal glucocorticoid metabolites. Under the first objective, the influences of some of the fundamental intrinsic and extrinsic factors on the fGCM levels were also assessed. 2. The second objective was to assess the association between body condition, seasonality and stress response. 3. Having assessed the relationship between body condition and stress-status, the third objective focused on the influence of socio-ecological correlates such as group size, lactational status and the presence of adult females in a herd on the stress status of female adult elephants. 4. With above three objectives majorly highlighting the influence of ecological factors, the fourth objective was to assess the influence of anthropogenic factors on stress-response of free-ranging elephants under which we compared the stress-response between crop-raiding in human-dominated and nonraiding elephants in forested habitats. The study was conducted during dry season (February to May) and wet season (August to December) of 2013 and 2015. The dissertation is organized in the following way: Chapter 1: General Introduction The first chapter provides a general introduction about stress response and its physiological pathway. It also discusses the importance of non-invasive technique used in our study. It highlights the significance of this study based on the available literature on both African and Asian elephants. Chapter 2: General validation and standardization of field techniques and assays; influence of intrinsic and extrinsic factors on faecal glucocorticoid metabolites This chapter is divided into two parts. The first part deals with general validation and standardization of field techniques and the second part assesses the influence of fundamental intrinsic and extrinsic factors. Being the first study on stress-response of free-ranging Asian elephants, influence of various field-based techniques and variations were studied to facilitate the reliable interpretation of fGCM levels. We first conducted the experiments to standardize the field protocol and the laboratory protocol. Under the field protocol, we found that there was no within sample variation (as observed in many other species) in fGCM levels from thoroughly mixed faecal samples and samples from the center of the same dung bolus. The time of collection of samples had a significant influence, suggesting that the concentration of fGCM remains stable until 6-9 hours. Influence of storage of lyophilized samples suggested that the concentrations of fGCM levels decline if exposed to moisture. Under the laboratory protocol, A group specific 11-oxoetiocholanolone EIA, used for analyzing fGCM levels (μg/g), was modified to enhance the repeatability, accuracy and precision. EIA was analytically validated in every hormonal analysis by calculating the inter- and intra-assay coefficients of variation, specificity and parallelism. In the second part, we assessed the influence of age, sex and season on fGCM levels and found there was no age-wise variation and sex-wise variation. Season-wise variation was significant only in female elephants. We discuss the potential physiological reasons behind such variation. These findings suggest that aspects such as time of collection, proper storage of samples and the influence of extrinsic stresses (seasonality) should be considered for reliable and unbiased interpretation of fGCM levels. Chapter 3: Assessment of season-dependent body condition scores in relation to faecal glucocorticoid metabolites in free-ranging Asian elephants. (Published in Conservation Physiology; Pokharel et al., 2017; doi:10.1093/conphys/cox039) We studied seasonal and annual changes in visual body condition scores (BCS), and assessed how these scores were related to levels of fGCM levels in free-ranging Asian elephants in the seasonally dry tropical forests of the Mysore and Nilgiri Elephant Reserves in southern India. We assessed the animals’ BCS visually on a scale of 1 to 5; where 1 represents a very thin and 5 represents a very fat elephant. To understand the influence of seasonality on BCS, we sampled the population during dry and wet seasons of 2013 and 2015 while, for annual changes in BCS, we sampled nine free-ranging adult females from different family groups that had been repeatedly sighted over seven years. To evaluate the influence of body condition on fGCM, we measured the fGCM levels from fresh fecal samples collected from the body-condition scored animals. Effect of age and season on BCS in relation to fGCM was also studied. We found that the BCS was related with season, i.e. individuals with low BCS were more frequent during the dry season as compared to the wet season and fGCM levels were negatively associated with BCS. To be precise, concentrations of fGCM were highest in individuals with the lowest BCS (BCS-1) and then significantly declined till BCS-3. fGCM levels were almost comparable for BCS 3, 4 and 5. This pattern was more conspicuous in female than in male elephants. We discussed the possible reasons behind such relationship between fGCM and BCS. The findings from this chapter suggest that season-dependent BCS, hence, reflect the stress status as measured by fGCM, especially in female Asian elephants and could be used as an important non-invasive approach to monitor the physiological health of free-ranging elephant populations. This is the first study to compare the body condition scores and stress-response in free-ranging elephants. Chapter 4: Influence of ecological and physiological correlates on stress physiology of free-ranging female Asian elephants. We investigated the influence of herd size (HS), lactational status and number of adult females present in a herd on adrenocortical function in free-ranging adult female Asian elephants by measuring their fecal glucocorticoid metabolites (fGCM) levels. We found that there was a negative association between herd size and fGCM levels when only HS and fGCM were compared. Lactating females showed higher fGCM than that observed in non-lactating females, which might reflect the nutritional stress on mother and anti-predatory challenges against the calves. Herds with only one adult female had higher fGCM levels than herds with more than one adult female which could be because of social challenges. The poor association between herd size and fGCM when all predictor variables were combined elucidate that group size may not necessarily influence the adreno-cortical function. However, the numbers of adult females present in a herd and their lactational status has a significant role in defining the stress-response in free-ranging adult female Asian elephants. This study is the first study to document the influence of lactational status with the stress-response in free-ranging elephants. Chapter 5: How physiologically costlier it is to be a crop-raider in a human-dominated landscape? Diet quality as a possible ‘pacifier’ against stress. We studied the stress-response in crop-raiding elephants in a human-dominated landscape and non-raiding elephants in the protected forested areas. While raiding agricultural crops, elephants face various associated threats such as retaliation by humans, human-induced disturbances and stress of raiding which could enhance the energetic costs, ultimately elevating their stress levels. We hypothesized that crop-raiders (in human-dominated landscape; Hassan district of Karnataka, India) will exhibit higher faecal glucocorticoid metabolite (a proxy of stress-response; fGCM) levels as compared to the nonraiders (in forested areas; Bandipur and Nagarahole National Parks). Contrary to our hypothesis, fGCM levels were found to be higher in both nonraiding female and male elephants than female and male crop-raiders. To assess the influence of benefits obtained from crop-raiding as one of the possible factors for the lower fGCM in crop-raiders, the difference in vegetation between human-dominated landscape and forested areas were further analyzed by using a remotely sensed NDVI (Normalized Differential Vegetation Index) and was ground-truthed by measuring the quality of diet (C:N ratio through fecal samples). Interestingly, the NDVI values were higher at the human-dominated and the C:N ratio was lower (higher Nitrogen content) in faecal samples of crop-raiding elephants (both in females and in males) than the nonraiders in the forested areas. Positive correlation was observed between C:N ratio and fGCM levels. These findings suggest that crop-raiding comes with the benefits of easy access to good quality of diet which may help in reducing the stress-response in elephants while being in the human-dominated landscapes, provided there is a low intensity of human disturbance. This is the first study to document and compare the stress-status of crop-raiding Asian elephants with elephants in their natural habitats using NDVI and C:N ratio. Chapter 6: Conclusion In conclusion, the work presented in this dissertation provides an overview of how ecological and anthropogenic factors could influence the physiological health of free-ranging Asian elephants. This dissertation focused on providing detailed insights about the stress-response in free-ranging Asian elephants. The objectives have been achieved by assessing the relationship between ecological stressors such as seasonality, body condition, herd size, lactational status, a presence of individuals in a herd and anthropogenic stressors such as human-induced threats or crop-raiding using well-validated and standardized laboratory and field protocols. This study provides valuable insights into the physiological health or stress-response being synergistically influenced by various ecological, social and anthropogenic factors in free-ranging Asian elephants. Findings obtained from this study could help in addressing the issues related to the management of free-ranging elephant populations.

The Asian elephant (Elephas maximus), a charismatic ‘flagship species’, is threatened by extinction in the wild, and the development of self-sustainable captive populations is a key conservation challenge. A third of the Asian elephant population is presently in captivity and information on the reproductive status, especially in females, is still lacking to a large extent. The onset of estrus in female Asian elephants is not associated with any visible physical signs, thus making the assessment of the reproductive status rather difficult. One approach to understanding reproductive cyclicity of animals is through generating profiles of reproductive hormones in blood (Wiseman et al. 1983; Brown et al. 1999, 2004). Profiles of reproductive hormones such as progesterone (P4), estrogen, luteinizing hormone and follicle stimulating hormone have already being demonstrated in Asian elephants (Brown et al. 1999, 2004; Brown 2000). In these studies, the reproductive status of females, maintained in zoos/captivity, was characterized based on circulating levels of hormones in blood samples. This is difficult to implement in the case of semi-captive or wild populations of elephants due to practical, legal and ethical considerations. In order to overcome this problem and to better understand the estrous status of female elephants, it is important to develop and validate non-invasive methods to monitor the reproductive status of female Asian elephants. An alternative approach to evaluating the reproductive status of females is to consider behavioral responses shown by males towards chemical signals produced by females to advertise their reproductive status. In order to understand the reproductive status of the individual belonging to the opposite sex, studies have shown that elephants rely on a variety of chemical signals produced in biological fluids such as urine, temporal gland secretion, inter-digital gland secretion, etc. (Krishnan 1972; Rasmussen & Schulte 1998). Chemical signaling is one of the prominent modes of communication in elephants, especially with respect to locating potential mates (Sukumar 2003). Thus, in most cases, elephants usually employ specific behavioral responses, for example sniff, check and place behaviors of trunk, for investigating the reproductive status of the conspecific individual, belonging either to the same or the opposite sex (Rasmussen et al. 1996; Schulte & Rasmussen 1999; Bagley et al. 2006). The objectives of the thesis are two-fold. First, to develop a non-invasive method of reproductive monitoring from fecal hormonal metabolites and also to understand the possible role of feces as an inter-sexual signal. The main body of thesis is divided into four chapters. 1) Development and validation of a non-invasive method to estimate progesterone metabolite in feces, to monitor the reproductive cyclicity of female elephants (Chapter 2). 2)Generation and characterization of progesterone and its metabolite, 5α-P-3-OH, profiles of semi-captive females using the developed non-invasive method to measure fecal metabolites (Chapter 3). 3) Validation of developed methodology and assay systems to a wild-population of female elephants (Chapter 4). 4) Feces as a potential source for inter-sexual chemical signaling in Asian elephants (Chapter 5). The above studies were carried out on semi-captive male and female elephants maintained in the forest camps of Mudumalai Wildlife Sanctuary (MWLS), Tamil Nadu and Bandipur National Park (BNP), Karnataka, India (Chapters 2, 3 and 5). For Chapter-4, free-ranging females of the MWLS were examined. 1. Development and validation of a non-invasive method to estimate progesterone metabolite in feces, to monitor the reproductive cyclicity of female elephants Niemuller et al. (1993) generated a profile of the progesterone metabolite, 5βpregnanetriol, to assess the estrous phase of Asian elephants based on non-invasive urine sampling. However, the collection of urine is difficult and to some extent impossible in the case of semi-captive and as well as that of wild elephants. Thus, the method of choice in this study was the development and validation of a non-invasive approach to measure fecal progesterone metabolites to assess reproductive status of females. Sampling was carried out at monthly intervals on three female elephants at the MWLS forest camp, while three other females maintained at the BNP forest camp were sampled fortnightly. An enzyme linked immuno-sorbent assay was developed to measure the concentration of the progesterone metabolite, 5α-P-3OH in the fecal samples of the semi-captive females. Using varying concentrations of the hapten (5α-P3OH), from low (0.1 mM) to high (1 mM), a standard curve was first generated, which had a linear range between 0.25 mM to 62.5 mM, with an EC50 of 1.37 mM. The linear range was then used to detect the concentrations of 5α-P-3OH in the fecal samples of females examined. The non-invasive method was further validated as there existed a positive correlation (p<0.1) between the levels of fecal 5α-P-3OH and that of concentration of circulating P4, measured in blood samples. This is the first valid documentation of a non-invasive method based on fecal progesterone metabolite pattern in order to assess the reproductive status of the female Asian elephants. 2. Generation and characterization of reproductive hormone profiles of semi-captive females using the developed non-invasive method to measure fecal metabolites With the establishment of a non-invasive method to understand occurrence of estrus in female elephants (described in Chapter 2), attempts were made to generate hormonal profiles over a longer time interval through more frequent sampling. Based on sampling at weekly intervals, the concentrations of both fecal 5α-P-3OH and that of native P4 hormone in the blood were determined, and the females were then identified as belonging to different reproductive states of hormonal cycling (follicular and luteal phases), non-cycling and pregnant categories. Of the 7 females sampled at both MWLS and BNP, three distinct categories of hormone-metabolite profiles emerged. The first category included four females that showed regular cyclicity throughout the sampling period, as monitored through the measurement of fecal 5α-P-3OH and serum P4 levels. The estrous cycle of all the four females was divided into two phases (follicular and luteal), based on the patterns of fecal 5α-P-3OH and serum P4 concentrations. The follicular phase of the estrous cycle was assigned when the values of both fecal 5α-P-3OH and serum P4 remained below 0.3 µg/gm and 0.3 ng/ml, respectively, for a considerable time period (viz. >2 wk). However, the luteal phase was characterized, when the levels of fecal 5α-P-3OH and serum P4 remained at or above 0.3 µg/gm and 0.3 ng/ml respectively, over a period of more than 2 - 3 wk. The second category had two females showing a ‘flat-lining pattern’ for the levels of fecal 5α-P-3OH and serum P4 concentrations, without any peak or dip in their concentrations. Since the reproductive pattern for both these females was flat-lined throughout their sampling period (51 wk), maintaining the levels of fecal 5α-P-3OH and serum P4 below 0.3 µg/gm and 0.3 ng/ml respectively, the females were considered to be non-cycling or anestrus. The third category had one female in which the levels of both fecal 5α-P-3OH and serum P4 measured were consistently high throughout the entire sampling period (26 wk). The levels of both fecal -P-3OH and serum P4 were above 0.3 µg/gm and 0.3 ng/ml, respectively. At the end of the sampling period, this female delivered a male calf; thus, the measured concentrations of fecal 5α-P-3OH and serum P4 signified the levels maintained during the gestational phase of this female. Sampling and hormonal analyses were also carried out for a male in the MWLS forest camps to investigate the baseline concentrations of fecal 5α-P-3OH and serum P4. The male showed consistently low concentrations of both fecal 5α-P-3OH and serum P4 throughout the sampling period (16 wk), below the margin of 0.3 µg/gm and 0.3 ng/ml, respectively. In this part of the study, it was confirmed that the reproductive status of a female elephant can be correctly assessed on the basis of measurements of fecal 5α-P-3OH alone, with repeated sampling of the female over a longer time scale. It was also shown that the strength of the positive correlation between the concentrations of the fecal 5α-P3OH and the serum P4 increased (p<0.01) for a larger sample size than that obtained for a comparatively smaller sample (described in Chapter 2). 3. Validation of developed methodology and assay systems to wild-population of female elephants So far, the non-invasive method to estimate fecal progesterone metabolite in order to predict occurrence of estrus of elephants was largely applied to semi-captive females (described in Chapters 2, 3). However, the necessity of such a method is being recognized for assessing the reproductive status of free-ranging females. Several findings have described differences in the rate and type of steroid metabolite excretion among individuals maintained under different diet regimes (Wasser et al. 1993; Smith et al. 2006). For instance, female elephants in the forest camps are provided with supplementary diet consisting of sugarcane, rice and millets. This diet is strikingly different from the feeding materials consumed by wild/free-ranging elephants (Sukumar 2003). Therefore, differences in dietary components can potentially influence the fecal steroid metabolites’ excretory patterns shown by semi-captive vis-a-vis wild females, which can affect the validity of measuring fecal 5α-P-3OH to predict females’ reproductive status. In order to examine this problem, the non-invasive method was applied to the population of wild elephants in the forests of MWLS, through random one time sampling of 30 individual female elephants. The steroid extraction efficiency (73 ± 11.0%, mean ± S.D., n =30) determined for the fecal samples collected from the wild females was not significantly different from the coefficient calculated in the case of the semi-captive females (80 ± 4.3%, mean ± S.D., n = 38). This indicated that dietary differences between wild and captive elephants did not influence levels of fecal hormonal metabolites in feces, unlike earlier observations on baboons (Wasser et al. 1993), old world primates (Wasser et al. 1988) and sheep (Smith et al. 2006). The values of the fecal allopregnanolone determined in the case of the wild females, ranged from as low as 0.06 µg/gm to as high as 23µg/gm of the sample, thus showing the heterogeneity of the samples, indicating that the females may be belonging to different reproductive phases. However, since sampling was carried out randomly, with an adult female being sampled just once, at this stage, it is not possible to identify or elaborate on the reproductive phase of the females. This is the first study reporting the values of the fecal progesterone metabolite in female Asian elephants in the wild. Further studies may be required to carry out long term monitoring of the wild females, through repeated collection of fecal samples over time from particular female. 4. Feces as a potential source for inter-sexual chemical signaling in Asian elephants In this Chapter-5, behavioral trials were conducted on male elephants to understand the role of fecal matter in conveying inter-sexual chemical signals. This was demonstrated by analyses of specific chemosensory behavioral responses shown by males towards the fecal samples of females that were strangers and belonging to different reproductive stages. Males showed four prominent behavioral responses namely ‘distant sniff’, ‘close sniff’, ‘check’ and ‘place’ towards the fecal samples of females. The sum of frequencies of these four responses (distant sniff, close sniff, check and place) was much higher for samples of the follicular (pre-ovulatory) phase females as compared to those of the luteal (post-ovulatory) phase females (p<0.005). Thus, for the first time, it was shown that male elephants are able to discriminate the different reproductive phases of females based on their specific behavioral response towards the fecal samples of the opposite sex. In conclusion, the thesis has focused on understanding and providing new insights regarding the reproductive biology of the female Asian elephants. This has been achieved through the development of the non-invasive method based on measuring the concentrations of the fecal progesterone metabolite and through the analyses of the chemosensory responses performed by the males towards the fecal samples of strange females. These methods can potentially be applied to the populations of both wild and captive/semi-captive female elephants in order to evaluate their reproductive status, through non-invasive measures. The information derived from the application of such methods will help in understanding the reproductive potential of the wild elephants under various environmental and ecological conditions. Further, the non-invasive measurement of reproductive hormones will help in monitoring the reproductive state of the individuals and thus aid in planning strategies for the welfare and management of the elephants maintained in captive or semi-captive conditions.

Dissertação de Mestrado Integrado em Medicina Veterinária, Ciências Veterinárias
Os elefantes Africanos e Asiáticos são considerados espécies ameaçadas e as suas populações continuam a decrescer. Inúmeros esforços têm sido feitos de forma a contrariar esta tendência. Os Zoos têm um papel muito importante no que toca à conservação da vida selvagem, desenvolvendo projectos de investigação e programas educacionais. Portanto, a existência de animais em vias de extinção em cativeiro é uma mais-valia para a sua sobrevivência. No entanto, estas instituições têm de mostrar ao público que a sua missão principal é a conservação de espécies, e que durante este processo todas as medidas com vista a garantir o bem-estar animal são acauteladas. O objectivo principal deste trabalho é avaliar as variações das manifestações comportamentais das diferentes populações de elefantes nos Zoos face às diferentes variáveis ambientais a que estão sujeitos. Para alcançar este objetivo, foram colhidos dados em cada Zoo sob a forma de questionários e de vídeos comportamentais de elefantes, que foram depois analisados estatisticamente. Observamos que a presença do macho influencia nas fêmeas os comportamentos de alimentação, locomoção, vocalização e comportamentos estereotipados. O comportamento de “dusting” e os movimentos das orelhas estão diretamente relacionados com o aumento da temperatura. Os dispositivos de enriquecimento levam a um aumento do comportamento de brincadeira, mantendo os animais mental e fisicamente ocupados. O enriquecimento alimentar que leva a um aumento do comportamento de alimentação é um factor positivo no que toca a comportamentos estereotipados, já que estes não são tantas vezes manifestados quando esse tipo de enriquecimento é praticado. Também pudemos observar que a existência de grupos sociais leva à realização de comportamentos mais semelhantes aos seus co específicos selvagens. Podemos concluir que a avaliação do comportamento dos elefantes em cativeiro permite auxiliar a adoptar medidas com vista a melhorarem os procedimentos de maneio em cativeiro, de forma a diminuir o impacto negativo na vida dos animais levando a uma melhor conservação.
African and Asian elephants are listed as threatened species and their populations continue to decline. Lots of efforts have been made to counter this tendency. Zoos have an important role to play in terms of Wildlife conservation, developing projects in the wild, alongside research and education programs. So, the existence of endangered species in captivity is important for their survival in the wild. Zoos must make clear to the general public that their mission is one of conservation, which is conducted with the highest welfare standards. The main objective of this project is to evaluate the behaviours’ variations in different Zoo populations regarding the effect of diverse environmental variables. To achieve this, in each Zoo was collect data through two methods: a survey and videos of elephant behaviours that were statistical analysed afterwards. We observed that the presence of the male has an influence on females’ foraging, locomotion, vocalizations and stereotypic behaviour. Dusting and ears’ flapping behaviours are directly related with the environmental temperature. The enrichment devices lead to an increase in the play behaviour, keeping the elephants mentally and physically occupied. The nutritional enrichment which leads to an increased the foraging behaviour is a positive factor regarding stereotypic behaviours as they are not performed as often as when such enrichment is practiced. Also, the existence of social tiers and groups leads to the performance of most similar behaviours as their wild conspecifics. We can conclude that the assessment of elephant behaviour in captivity will help to adopt measures to improve the management procedures that will have lower impact in the animal’s life leading an optimized conservation.