David Anderson and Mary Wood, University of California, Davis - Friday, 10:00
Web-based Search Templates on Elephants

This presentation introduces a web-based gateway, providing convenient and user-friendly access to information resources and bibliographic search templates concerning elephants. This gateway, Gateway to Information on Elephants, offers direct links to some outstanding bibliographies, databases, and articles on elephants. It also links directly to various databases associated with the California Digital Library and includes some embedded searches, "search templates", that make it possible to conduct new searches on elephants with previously stored search instructions. Among the various databases that are linked, some are available to the general public, some only to UC affiliates, and some to UC Davis affiliates. The section on searches includes the specific key words that are especially useful within each databases, since the conventional terminology varies somewhat across databases. http://www.vetmed.ucdavis.edu/CCAB/elepha~1.htm
 

Robert H. I. Dale, Nicole Jordan, Sarah Kinnett, Lindsay Beach, and Jenny Noble, Butler University, Indianapolis, Indiana - Friday, 3:20
Behavioral Development of Elephant Calves: Review with Examples from the Indianapolis Zoo

Many observers have described aspects of the development of elephant calves, for both Asian elephants (Elephas maximus) and African elephants (Loxodonta africana). We have drawn from a variety of sources to produce a generic description of the behavioral development of the elephant calf. We have attempted to identify developmental milestones for physical growth, sensori-motor abilities, nonsocial behaviors (such as self-grooming), and social behaviors (e.g., play). Our primary goal was to integrate the contributions of our colleagues, rather than to identify new behaviors ourselves. We present data from wild elephants, captive herds, and "tame" elephants (wild-captured, actively managed Asian elephants). A secondary goal of the paper is to address the behaviors of captive elephants, especially those of calves. We describe the similarity of the behavior of captive calves to the behavior reported for calves in the wild herds. Our preliminary data suggest that the calves at the Indianapolis Zoo are following the same developmental schedule, and exhibiting the same behavioral repertoires, as calves in wild herds. We will provide photographic and video examples of some of these behaviors.
 

Karen Emanuelson, Oakland Zoo - Saturday, 11:50
Protected Contact and Medical Care in Captive Elephants, with a Case Presentation of Salmonellosis in Elephants at the Oakland Zoo

The Oakland Zoo has managed four African elephants, a bull and three cows, in a Protected Contact system since 1991. The types of behaviors trained at the zoo, with an emphasis on medical behaviors, will be discussed. A severe outbreak of Salmonellosis that occurred in the elephants, resulting in two abortions, will be described. The administration of intensive medical care in a Protected Contact system will be discussed.
 

Laurie J. Gage, David Blasko and the Elephant Staff. Six Flags Marine World - Saturday, 11:20
Husbandry and Medical Considerations for Geriatric Elephants

Within the next decade, over half of the existing population of Asian elephants in north America will be over 40 years of age. Presently over 60% of Asian elephants in north America listed with ISIS are already over the age of 30. And in the next decade about 50% of the existing captive population of African elephants will be 30 years or older.
Foot problems have posed major medical problems for many of the older captive elephants. Osteomyelitis in the bones of the feet of older elephants has been a significant cause of mortality in the over 40-year age group. The best method of treating osteomyelitis is to take preventive measures to keep the infection from occurring in the first place. Preventing foot problems has been a key topic for elephant veterinarians, trainers, and keepers. Anecdotal and scientific information point to several husbandry and veterinary factors that may contribute to resolving the problem. These include:

Exercise: Elephants that have significant daily exercise seem to have fewer foot problems than elephants that primarily stand throughout the day. We have found that regular daily exercise is extremely important, and older elephants that were retired and not encouraged to walk each day developed toe and foot infections at a greater rate than their active counterparts.

Type of substrate or footing: Prolonged housing on hard surfaces such as concrete or asphalt should be avoided, and the use of dirt, wood or rubberized footing in primary enclosures should be the standard. We believe that elephants spending the majority of their lives on hard surfaces such as concrete or asphalt are more prone to foot problems than elephants housed on softer or more natural substrates. We feel concrete floors in primary housing areas should be covered with a substrate with more give to it, such as a rubberized coating, thick rubber mats, or wood.

Diet : Captive elephants are prone to deficiencies, especially vitamin E, and should receive a daily supplement of the vitamin either in the form of the water-soluble tocopherol, or the natural vitamin E found in fresh browse provided daily. Elephants should have blood drawn to screen them for baseline circulating levels of vitamin E to ensure their supplementation is adequate. Biotin is another supplement suggested to be necessary in the diet to maintain the health of the feet and nails. Elephants should be fed a clean high quality diet, however, it is important to feed hay or browse that is lower in caloric value than diets fed to domestic animals. Elephants that are overweight may be more prone to foot problems than other elephants.

Regular foot care by skilled personnel: Probably one of the most important methods of preventing foot or toe infections is an aggressive foot care program. Feet of all captive elephants should be examined regularly, and elephants in both free or protected contact should be trained to allow regular foot examinations and sole and nail care. Maintaining nails and soles is supremely important in the prevention of foot infections. Frequent sole and nail examinations and regular trimming of nails and soles is a must for captive elephants, especially for those that do not receive adequate daily exercise.

General health considerations: The overall health of an elephant may also contribute to the health of its feet. Elephants with illnesses or metabolic disorders may be more prone to foot infections, and may not be able to mount an appropriate immune response to a small foot infection, thus enhancing the chance of the infection progressing into the bones of the foot, causing osteomyelitis. Treating systemic medical problems aggressively and supporting the immune system are important to the overall health of the elephant, and important to the health of their feet.

Preventive care or treatment of arthritis in older elephants: Treating arthritic conditions with non steroidal anti-inflammatory drugs, low level infra-red light therapy, polysulfated glucosaminoglycan (Adequan), hyaluronate sodium injections or other supportive care therapies will increase the comfort of the arthritic elephant and enable the animal to exercise more frequently without discomfort. The regular exercise in turn seems to improve the general health of the feet.

Veterinary preventive foot care regime: Survey foot radiographs are important to obtain before problems occur, as they offer a comparison of the normal foot to the diseased foot. The use of appropriate topical treatments, patches, shoes or pads are important measures to prevent minor foot problems from progressing.

Aggressive veterinary treatment of new foot infections: This includes regular radiographic foot studies, early culture and sensitivities when a lesion is first discovered, and appropriate systemic or localized antibiotic therapy.

In conclusion, older elephants may have greater need for supplements and routine exercise to help to prevent foot and other health problems. Housing elephants from an early age on appropriate substrates will likely mitigate foot problems in the future.
 
 

Thomas E. Goodwin, Chemistry, Hendrix College, Conway, Arkansas - Friday 1:00
The Secrets in Secretions: Unraveling Elephant Mysteries via Chemical Methodologies

Behavioral observations of elephants have long led to speculation that a primary function of temporal gland secretions (TGS) is chemical communication (Sikes, 1971). This function has been clearly demonstrated in recent years for Asian elephants through chemical analyses and bioassays by Professor Bets Rasmussen and her co-workers (Rasmussen, 1998). A seminal study of African elephant TGS identified three unexpected sesquiterpene constituents: farnesol, a well-known natural product, and two novel farnesol derivatives (Wheeler et al., 1982). Limited bioassays of these compounds were inconclusive (Gorman, 1986; Rasmussen, 1988).

In collaboration with Professor Rasmussen, we have initiated a new investigation of temporal gland secretions from African elephants in order to identify trace organic chemicals dissolved therein. The impetus to pursue further analysis arose from the realization that much is yet unknown in the chemistry that may connect behavior, physiology, and chemical signaling. Our early studies confirmed the presence of the three sesquiterpenes mentioned above. In addition, we identified (E)-2,3-dihydrofarnesol, a bumblebee pheromone not seen before in mammals, and a rare component of a Greek tobacco, drimane-8?,11-diol, never observed before in an animal (Goodwin et al., 1999). In recent work as yet unpublished, two new sesquiterpenes, both farnesol analogues, have been identified in TGS from a number of widely dispersed African elephants. These are new chemical compounds that have never been reported in the literature from any source, natural or synthetic.

In this lecture, we will outline our methods of TGS collection, and the harvesting of trace organics therefrom using the relatively new technique of solid phase microextraction (SPME). The use of gas chromatography (GC) for separation of the mixture of components, and mass spectrometry (MS) for molecular structural assignment of novel compounds will be explained. Since thorough, multiple bioassays require larger sample sizes than can be obtained from TGS and these novel compounds are not available commercially, we have prepared them in the laboratory. These preparations also make possible the confirmation of structural assignments by comparison of natural and synthetic compounds. Finally, our methods for field bioassays will be briefly described.

Despite possible appearances to the contrary, the primary objective of this talk is not to overwhelm or confuse the non-specialist with arcane chemical details. We hope, rather, to provide a comprehensible overview of how the instruments and methodologies of organic chemistry can complement other methods of elephant study. These techniques can play a major role in the definitive delineation of temporal gland function, and will increase our understanding of chemical signals that govern elephant behavior.

References:
Goodwin, T.E., E.L. Rasmussen, A.C. Guinn, S.S. McKelvey, R. Gunawardena, S.W. Riddle, H.S. Riddle (1999) African Elephant Sesquiterpenes. J. Nat. Prod. 62: 1570-1572.

Gorman, M.L. (1986) The secretion of the temporal gland of the African elephant Loxodonta africana as an elephant repellent. J. Trop. Ecol. 2: 187-190.

Rasmussen, L.E.L. (1998) Chemical communication: An integral part of functional Asian elephant (Elephas maximus) society. Ecoscience 5: 410-426.

Rasmussen, L.E.L. (1988) Chemosensory responses to two species of elephants to constituents of temporal gland secretion and musth urine. J. Chem. Ecol. 14: 1687-1711.

Sikes, S.K. (1971) The Natural History of the African Elephant. American Elsevier, New York, pp. 34-38.

Wheeler, J.W., L.E. Rasmussen, F. Ayorinde, I.O. Buss, G.L. Smuts (1982) Constituents of Temporal Gland Secretion of the African Elephant, Loxodonta africana. J. Chem. Ecol. 8: 821-835.
 

B. L. Hart, L. A. Hart, M. McCoy, and C. R. Sarath; University of California, Davis; Jungle Lodges and Resorts, Nagarhole National Park, India - Friday, 3:50
Tool Use as a Marker of Cognitive Behavior in Elephants

Large-brained, long-lived mammals have captured the attention of scientists interested in animal cognitive functions. Various aspects of complex social interactions and communication have typified the studies of the great apes and cetaceans. Recently, tool use has emerged as a marker of cognitive behavior in some great apes and systematic studies have focused on the details of tool use and modification.

The terrestrial animals with the largest brains and that are arguably the longest living, namely Asian and African elephants, have been neglected relative to apes and cetaceans with regard to cognitive behavior. Yet, a historical note in 1838 by the African adventurer, W.C. Harris, records perhaps the first written account of tool use by any non-hominoid species. In one of his outings, Harris encountered elephants emerging into an open glade and states the elephants were " bearing in their trunks the branches of trees with which they indolently protected themselves from flies."

The studies reported here focus on fly switching as a prominent example of tool use in Asian elephants. We recorded several instances during the fly season where wild elephants used branches, ostensibly to repel flies while they were foraging. Systematic experiments on tool use utilized groups of captive elephants maintained in naturalistic or enriched environments in or near national parks. One study showed that fly switching frequency significantly correlated with the intensity of flies on or around the elephants. Another study found that fly switching significantly reduced the number of flies on or around elephants. A third study on captive elephants tested the animals' readiness to modify branches that were presented to them but which were too long or clumsy to use without modification. Most elephants (8 of 13) modified the branch by removing a side stem or distal section of the main stem and switching with this modified branch.

Several opportunistic observations have been made on wild and captive Asian and African elephants using tools, including, throwing sticks or rocks at other animals while foraging and scratching with a stick. Fly switching, when flies are prominent, may represent the most frequent form of tool use and the best example of tool modification in elephants.
 

Lynette A. Hart, University of California, Davis and Sundar; Jungle Lodges and Resorts, Nagarhole National Park - Friday, 9:10
The Mahout-Elephant Relationship: Ancient and Modern Versions

Elephants in India have long been used as working animals, especially as a means of Indian transport. This tradition of domesticating elephants featured a lifelong and
full-time close relationship between the mahout elephant-handlers and elephants. This presentation concerns how the mahout-elephant relationship has changed over time.

In early times, the elephants were owned by royalty and resided in a palace or at royal hunting grounds where the elephant and mahout lived as a unit. The life of a mahout was customarily passed from father to son. Often a mahout grew up and ultimately aged with his elephant.

With the advent of modern governments and the reduction of use of elephants in logging, elephants now are subject to private ownership (tourist industry) or governmental departments of forestry. The traditional lifestyle of domesticated elephants and their mahouts is rapidly disappearing and may soon be extinct. New roles for elephants such as giving rides to tourists or performing in temples, processionals, and festivals still provide some employment for elephants and the successors to traditional mahouts. These new mahouts, basically hired for the job of managing an elephant, still express respect and thoughtful consideration of their relationships with their elephants. Related counterparts to the mahout lifestyle can be seen in lifelong, full-time circus elephant trainers in the United States. Emerging practices for domesticating African elephants can be seen in Southern Africa, where the entire group of trainers works with all the elephants rather than developing a special relationship with a particular elephant.
 
 

Wendy Koch, U.S. Department of Agriculture - Friday, 10:30
Elephant Management Considerations from a Regulatory Perspective

Elephant management considerations from a regulatory perspective must take into account the concerns of the "clients" of the agency-activist and industry (and animal) interests-and at least two distinctive management categories- fixed vs. traveling displays (and the subcategories). Regulators must also deal with bureaucratic inertia, which greatly slows any needed regulatory changes. In an imperfect world, regulations are a compromise between science and politics and must balance reality vs. the ideal, especially with regard to ensuring safety of both animals and people. This talk will discuss current elephant management issues as affected by current regulatory requirements.
 

Kim Luikart, University of California, Davis - Saturday, 9:30
Anatomy of the Elephant Forefoot

This presentation will involve a laboratory demonstration of a prosected elephant leg, as well as radiographs, CT scans, and transverse slices of an elephant foot. The anatomy of the elephant foot is somewhat unique among animals. Asian elephants generally have five toenails in the front feet and four in back, while Africans usually have four toenails in front and three in back. This may vary between individuals and subspecies. The location of the digital bones is not readily apparent from external contours, although the position of the toenails may approximate the position of P3 in the corresponding digit. The palmar surface is covered with a keratinized pad, and a fatty digital cushion lies deep to the pad, interacting with the digits to provide an excellent cushion to support the great weight of each leg. There are 8 carpal bones which are arranged in two rows. There are 5 metacarpal bones, as well as 5 digits, and the stance of the elephant is digitigrade. In general, only digits 2, 3, and 4 contain 3 phalanges each. Digit 1 usually consists of only 1 phalanx, while digit 5 generally consists of 2 phalanges. The distal phalanges of digits 2-4 are attached to the dermis of the nail and are separated from the middle phalanges by connective tissue. The proximal sesamoid bones occur in pairs on the palmar aspect of metacarpals 2-5, and a single sesamoid is found on metacarpal 1. Elephants also possess a prepollex, a cartilaginous structure extending from C1 and MC1 into the digital cushion. This structure is assumed to aid in stabilizing the carpus over the digital cushion.

The primary blood supply to the front foot is from the median artery, which runs along the palmar aspect of the carpus and continues distal to the carpus as the metacarpal artery. The metacarpal artery branches palmar to the metacarpal bones, forming the first dorsal digital artery, then arches to form the deep palmar arch. The metacarpal arteries for digits 2-5 branch from this arch, becoming the palmar digital arteries and the dorsal metacarpal arteries, which continue on as the dorsal digital arteries for digits 2-5. The digital cushion is supplied by the palmar digital arteries.

The primary nerves of the front foot consist of the radial, ulnar, and median nerves. The radial nerve supplies the dorsal digital nerves for digits 1-3, while the dorsal branch of the ulnar nerve, which joins the radial nerve, supplies the dorsal digital nerves for digits 4 and 5. The median nerve supplies the palmar digital nerves for digits 1-4, whereas the palmar branch of the ulnar nerve forms the palmar digital nerve of digit 5. The digital cushion is supplied by the palmar digital nerves.

Few precise anatomical details exist in the literature regarding elephant foot anatomy, especially from a clinical perspective. The most common site of medical problems observed in elephants at zoological institutions is the feet. Cracks and abscesses in the nail, cuticle, and sole are common and often result in deeper infections. These infections spread to the bones and joints of the feet, resulting in osteomyelitis and arthritis. This anatomical study was undertaken in hopes of improving knowledge about the osteologic, vascular, and neurological structures of the foot, especially in relation to the development and persistence of chronic foot lesions. Increased knowledge of these anatomic features may help in the treatment and prevention of chronic pododermatitis. We have encountered abnormal digit conformation associated with chronic sole lesions in both specimens studied thus far, although more research must be done to validate this association. More studies are needed to improve the foot health of captive elephants, as well as to elucidate new possibilities such as seismic propagation through mechanoreceptors in the feet or toes.
 

Brenda McCowan, University of California, Davis - Saturday, 8:30
Developing a Quantitative Method for Analyzing Infrasonic Vocalizations in Elephants

Elephants produce infrasonic vocalizations that are thought to function in coordinating group movement across long distances. These vocalizations occur at frequencies below the range of human hearing and thus pose a particular challenge for developing a method that can sufficiently capture the salient variation likely inherent in these vocalizations. New analytical techniques are needed to capture this acoustic variability, and thus the potential use of these vocalizations in individual identification, group membership, and different contexts. Low frequency signals are more constrained in the amount of absolute frequency modulation than signals of higher frequency. Thus infrasonic vocalizations pose a particular challenge in deciphering which acoustic parameters appropriately capture salient acoustic variation.

I will discuss how modern computer techniques can be used to begin to unravel the acoustic structure and variation in elephant infrasonic vocalizations. Using a few examples, I will demonstrate and discuss how relative frequency modulation among other frequency, temporal and amplitude parameters can be measured from elephant vocalizations and eventually correlated with individual identity, group membership and different behavioral contexts.

Relevant publications:
McCowan, B. (1995). A new quantitative technique for categorizing whistles using simulated signals and whistles from captive bottlenose dolphins (Delphindae Tursiops truncatus). Ethology 100: 177-193.

McCowan, B. and Reiss, D. (1995). Quantitative comparison of whistle repertoires from captive adult bottlenose dolphins (Delphindae Tursiops truncatus): a re-evaluation of the signature whistle hypothesis.  Ethology 100: 193-209.

McCowan, B. and Reiss, D. (1995). Whistle contour development in captive-born infant bottlenose dolphins: role of learning. Journal of Comparative Psychology 109 (3): 242-260.

McCowan, B., Reiss, D. and Gubbins, C. M. (1998). Social familiarity influences whistle acoustic structure in adult female bottlenose dolphins (Tursiops truncatus). Aquatic Mammals 24: 21-40.

McCowan, B., Hanser, S. F. and Doyle, L. R. (1999). Quantitative tools for comparing animal communication systems: information theory applied to bottlenose dolphin whistle repertoires. Animal Behaviour 57: 409-419.
 
 

Susan K. Mikota and Hank Hammatt, Sumatran Elephant Healthcare and Conservation Program -
Saturday, 2:10
Sumatran Elephant Healthcare and Conservation Program

Introduction
As human population growth soars past 6 billion, habitat for nature continues to diminish. Protected areas have become critical refuges for many animals and plants. With few exceptions, there is no "wild." Free-ranging and captive populations of endangered species must be cohesively managed to optimize limited human and monetary conservation resources. The combined efforts of scientific research, animal welfare initiatives, and financial commitments from private and governmental agencies will be needed to save many species.

Indonesia is the fourth most populous nation in the world. Sumatra, Kalimantan, Sulawesi, and Irian Jaya have absorbed over 2.5 million people as part of a government repatriation to relieve overcrowding on Java and other islands. There are plans to relocate an additional 65 million people over the next 20 years.

History of Elephant Conservation Centers (ECC's) on Sumatra
As a result of increasing human-elephant conflict, the government of Indonesia began establishing Elephant Training Centers (ETC's) on Sumatra in 1986. It was intended that the ETC's would train "problem" elephants for use in logging, patrol work, and tourism. Today there are six ETC's (now referred to as Elephant Conservation Centers or ECC's) that hold approximately 400 elephants. The Sumatran elephant is the most endangered Asian elephant. The elephants in captivity are a significant portion of the total population and may be critical to its survival.

Surveys conducted by the World Wildlife Fund (WWF), Fauna and Flora International (FFI), and others have been in agreement regarding the healthcare and husbandry problems of captive Sumatran elephants. Elephants suffer from parasites, marginal nutrition, and wounds. Some health problems are a direct result of training techniques. Most ECC veterinarians receive little or no elephant training. They have few resources for medical information and limited veterinary supplies. As a result, they are often poorly motivated and serve only brief terms. Economic crises in Indonesia and the current transition to a de-centralized government have seriously limited financial resources available to the centers. The anticipated use of elephants in logging has not materialized, and there are few revenues from tourism or patrol work. The long-term welfare of the Sumatran elephants will depend on external funding together with guidance from experienced elephant veterinarians and handlers. International communication among supporters would be enhanced by an in-country presence.

Sebanga-Duri Elephant Conservation Center
The Sebanga-Duri Elephant Conservation Center was established in Riau Province in east central Sumatra in 1988. It is located 115 km northeast of Pekanbaru (population of 700,000) and about 30 km southeast of Duri, a Caltex oil company community. A local decree designated 5000 hectares (~12,500 acres) for the Center in 1992. Villagers burned down the Center in 1993 and it was rebuilt in 1994. Squatters on ECC land have established three villages. Illegal logging and farming of oil palms have reduced the elephant preserve to fewer than 800 hectares. In January 1996, twelve local wild elephants were poisoned and buried. Law enforcement to prevent illegal settlements and logging does not seem to exist.

The monthly ECC budget for food is about 300,000 rp / elephant (~ $40 US). The monthly veterinary supply budget is less than one U.S. dollar per elephant, and even this is often used to purchase food rather than veterinary supplies. The current mahout salary is 250,000 rp / month (~ $ 35 US).

In April 2000, with support from Caltex, the World Wildlife Fund, and Fauna and Flora International, we conducted a medical evaluation of 41 of the 57 Sebanga-Duri elephants. Over a 7-day period, we established medical records, administered tetanus vaccinations and de-worming medications, collected blood samples, and initiated treatment for current health problems. Water used by the elephants for drinking was evaluated in response to questions raised in previous reports. We collected blood and tissue samples for the US Fish and Wildlife Service DNA databank.

A number of health concerns were revealed by physical examination and bloodwork. The internal parasite load is heavy. However, there is no routine worming program and no money for de-worming medications. Veterinary care is sporadic and supplies are grossly inadequate. Over 50% of the elephants are anemic. The quality of the diet is questionable and requires further study. Toxic levels of lead and cadmium were found in the drinking water and elephants may be suffering from chronic lead toxicity. Tuberculosis is endemic in the human population in Southeast Asia and captive elephants are at risk.

References
Krishnamurthy. V. 1992. Recommendation for improving the management of captive elephants in Way Kambas National Park, Lampung, Sumatra, Indonesia.
Gajah 9: 4-13.

Lewis, J. 1998. A veterinary assessment of Sumatran elephant training centers; a report on the visit of Dr. John Lewis of International Zoo Veterinary Group, on behalf of Fauna & Flora International, to the Sumatran elephant training centers at Lhokseumawe, Sebanga, and Way Kambas, Sumatra 29.04.98 - 13.05.98.

Lilley, R. and C. Saleh. 1998. Captive elephants in crisis; a WWF report on a survey of elephant training centers in Sumatra, Indonesia, 9-20 November, 1998. Submitted to WWF Asian Elephant Action Planning Workshop, Vietnam: 1-6 December, 1998.

Reilly, J. and P. Sukatmoko. 1998. The elephant training centre at Way Kambas National Park, Sumatra; a review of the centres operations and recommendations for the future. Dept. of Biological Sciences, Manchester Metropolitan University, PhD thesis.

Stremme, C. 1998. Significant veterinary problems caused by the training methods utilized by elephant training (ETC) in Sumatra, Indonesia; a report on behalf of Fauna & Flora International, to the Sumatran elephant training centers at Lhokseumawe, Sebanga, and Way Kambas, Sumatra 29.04.98 - 13.05.98
 

Caitlin O'Connell-Rodwell, Byron Arnason, and Lynette Hart, Stanford University, Tezar Inc., and University of California, Davis - Saturday, 9:00
The Seismic Propagation of Elephant Low Frequency Vocalizations and Possible Detection Mechanisms

As an elephant emits a low frequency vocalization, an exact replica of this signal is propagated in the ground. Based on our simultaneous acoustic and seismic recordings of elephant rumbles and using seismic modeling, we estimated that the elephant rumbles could potentially travel up to 16.7 km in the ground. Similar analyses of recorded foot stomps from a mock charge indicated a possible propagation of up to 32 kilometers. In an effort to determine if elephants can detect the seismic signals that they produce, we conducted seismic playback experiments with captive African and Asian elephants. Previously recorded seismic components of particular elephant vocalizations were played back through two specially adapted 50 W seismic transmitters at 10 m distances from the elephant. A TEAC digital recorder provided the signal source for the transmitters. Playback signals and elephant responses were recorded close to the elephant on a TASCAM 2-channel digital recorder through a 10 Hz Mandrel geophone and Neumann low frequency microphone. These signals were monitored real time through a Toshiba Satellite Laptop using SpectraPLUS software to ensure both signal integrity and that no seismic signals were coupling with the air. Detection of seismic signals was determined by a change in elephant behavior at the onset of the playback signals through video and visual observations. In addition, we are investigating anatomical specializations of the elephant's head and feet that could relate to seismic sensitivity as well as their low frequency auditory capabilities.

If seismic components of elephant vocalizations are detected by elephants, these animals may be communicating at much greater distances than previously thought. Seismic transmission would offer some advantages for long distance communication, since, unlike airborne transmission, it is relatively unaffected by environmental influences. Seismic vibrations received in combination with acoustic signals could be an important means of discriminating distance by judging the time lag between the arrival of the seismic versus the air borne signal. Determining the direction of the signal source might also be facilitated by the distance between an elephant's feet being greater than that between the ears, providing a feasible phase difference to distinguish between the times of arrivals of the acoustic and seismic signals. The results of this study may lead to a better understanding of elephant audition and long distance communication. The results of this study also raise the question of whether other large mammals might capitalize on available inter and intraspecific seismic signals for group cohesion, prey detection, or predator evasion.
 

Professor L. E. L. Rasmussen, Department of Biochemistry & Molecular Biology, Oregon Graduate Institute, Beaverton, Oregon - Friday, 1:30
Wild and Non-wild Elephants: How Two Modes of Olfaction and a Multitude of Chemical Signals and Pheromones Influence Elephant Behavior

Chemical signals in the Asian and African elephant are reflections of specific physiological and endocrinological events. Several identified chemical signals have been linked to discrete behavioral and/or endocrinological events; such linkage allows increased understanding of how elephants make their society functional. Recently, years of research on the influences of chemical signals among captive Asian elephants, Elephas maximus, has been confirmed among their wild counterparts and is being translated into knowledge useful for the preservation of the species (Rasmussen et al., 1996,1997; Rasmussen & Krishnamurthy, 2000).

The use of chemical signals among African elephants has been studied less and thus is less well understood (Goodwin et al., 1999). Nonetheless, field behavioral observations in Samburu of musth and nonmusth male African elephants clearly demonstrate the linkage between behavior and deposited chemical signals (Rasmussen & Wittemyer, in preparation).

Elephants of both species, with varying degrees of emphasis, utilize chemical signals from almost immediately at birth, if not before. Elephants are equipped with olfactory apparati unsurpassed in the mammalian world. For millennia, dating from written records from ancient India, the ability of elephants to precisely sense odors, often from significant distances, has been known. Recent studies with the pheromone, Z-7-dodecenyl acetate, have confirmed the exquisite sensitivity and precision of detection in the vomeronasal organ of adult male Asian elephants.

Some of the behavioral constraints imposed on young elephants by adults are regulated by chemical signals. The examples that will be discussed involve maternal-offspring recognition, the onset of estrous cyclicity in young females and "moda" musth in young males (Rasmussen & Riddle, in preparation).

By adulthood, batteries of chemical signals are present, some are single compounds and others complex mixtures. Now, molecular biology enables the study of functionality without injury to these endangered mammals, and analytical methods reach into detection limits of parts per trillion. In the captive adult Asian elephant we have demonstrated precisely what some of these signals are, the degree to which their content is apparently influenced by hormones, and what behaviors are linked to specific chemical signals. The most definitive chemical signal described so far is the female-dispersed urinary preovulatory pheromone, Z-7-dodecenyl acetate. Recent identification of Z-7-dodecenyl acetate in urine samples collected in the field from wild female Asian elephants, subsequent to observations of multiple flehmen responses by male elephants, is indeed exciting.

Our state of knowledge of other behavioral/chemical linked aspects of how elephants govern their own society is just at the frontier. Female Asian elephants tail flick at very specific times during their estrous cycle (Slade, 1999). To whom are they flicking their chemical signals? Field behavioral observations in collaboration with Dr. Krishnamurthy at Southern India sites suggest perhaps these tail-hair-dispersed signals wafting into the air are directed toward other females. The implications of such a speculation are discussed. Adult Asian males in musth, whether in captivity in USA or work camps in India or in the wild in India, emit frontalin, (which is also a bark beetle pheromone), large amounts of 2-nonanone, and a medley of other ketones, including cyclohexanone, in their temporal gland secretion, breath and urine (Perrin & Rasmussen, 1999; Rasmussen, 1998, Rasmussen, Krishnamurthy and Sukumar, in preparation). The chemical message from these mature males is very different from the sweet-smelling message of juvenile musth.

References:
Goodwin, T.E., Riddle, H.S., Riddle, S.W.,Guinn, A.C., McKelvey, S.S. & Rasmussen, L.E.L. (1999). African Elephant Sesquiterpenes J. Natural Products 62:1570-72.

Rasmussen, L.E.L. & V. Krishnamurthy (2000). How Chemical Signals Integrate Asian Elephant Society: The Known and the Unknown. Zoo Biology (in press).

Rasmussen, L.E.L. (1999). Elephant Olfaction ChemoSenses 2:4-5.

Rasmussen, L.E.L. & Perrin, T.E. (1999). Physiological correlates of musth: lipid metabolites and chemosignal composition. Physiology & Behavior 67: 539-549.

Rasmussen, L.E.L., Lazar, J., Greenwood, D., Feng, L., Prestwich, G.D. (1998). Initial characterizations of secreted proteins from Asian elephants that bind the sex pheromone, (Z)-7- dodecenyl acetate. Chemical Senses 23:591.

Rasmussen, L.E.L. (1998.) Chemical Communication: An Integral Part of Functional Asian Elephant (Elephas maximus) society. (1998) Ecoscience 5: 410-426.

Rasmussen, L.E.L., T.D. Lee, A. Zhang, W.L. Roelofs, & G.D. Daves, Jr. (1997.) Purification, Identification, Concentration & Bioactivity of (Z)-7-dodecen-1-yl acetate: Sex Pheromone of the Female Asian Elephant, Elephas maximus. Chemical Senses 22:417-438.

Rasmussen, L.E.L., Lee, Terry D., Roelofs, Wendell L., Zhang, A., & Daves, G. Doyle. (1996). Insect Pheromone in Elephants. Nature 379:684.
 
 

Jo-Ann Shelton, University of California, Santa Barbara - Friday, 8:40
Dancing and Dying: The Display of Elephants in Ancient Roman Arenas

In the western world, the use of elephants for entertainment may be traced to ancient Roman society. The purpose of this presentation is to discuss the types of spectacles that the Romans developed and the reasons why they were popular. The spectacles can be divided into two general categories: those in which elephants were mutilated and killed, and those in which elephants were made to perform stunts. To modern sensibilities, these categories seem quite different, the former involving death of an animal, and the latter involving a display of its talents. However, Roman spectators did not make the same distinctions. For them, the function of the events was to amaze and amuse, and they were equally amused by the tricks of a trained performer and the "tricks" which an injured elephant might employ to escape its tormentors. Elephant acts were popular because this species had been endowed with symbolic significance. For example, in the 3rd century BC, the elephant was identified with Rome's most hated military opponents, the Carthaginians of North Africa. Elephants were native to North Africa and were used as war machines by the Carthaginians. Thus, the torment of elephants in Roman arenas represented a victory over the defeated enemy, and it enabled the spectators to participate in the process of imposing Roman justice on a barbarian world. In addition, as the Romans expanded their power and territory, the display of elephants signified the success of their imperial policies. The capture and transport of elephants entailed enormous expense. The apparent ease with which the Romans were able to bring them to arenas for purposes of amusement offered proof that their state was powerful and prosperous. Of course, long before they had encountered elephants, the Romans had already developed spectacles in which animals were victimized. These spectacles provided reassurance that the orderly civilization that they had created could confront a hostile and irrational Nature and subdue it. As the largest land mammal, the elephant was an ideal candidate for the type of degradation and torment that seemed to validate human claims of superiority. Elephants were also ideal representatives of the victims of Roman political and military ascendancy. Their enormous size and strength made them appear menacing, but their lumbering gait and strange appearance meant that they could be easily ridiculed. Whether they were being forced to dress in human clothes and perform comical dances, or being slaughtered with javelins, elephants provided for Roman spectators a pleasure derived from knowing that they could bend the world to their will.
 
 

Patricia Simonet, Sierra Nevada College and Ramesh Krishnamurthy, University of the Pacific -
Saturday, 10:20
Self-recognition Among Captive Asian Elephants: Preliminary Results and Future Studies

One of the topics relevant to both cognitive science and animal behavior has been mirror self-recognition. This study examines mirror use and self-recognition in two Asian elephants, ages 8 and >45 years, for a period of two and one half months. During this period elephants used the mirror as a tool to find hidden objects, and to locate and investigate marks on their own bodies by way of the mirror. Observations were video recorded and logged in behavioral notebooks. Over 2,300 records contain detailed descriptions of sensorimotor, social, and communicative behavior. Two empirical observations during mirror studies have been interpreted as evidence of self-recognition: 1) when first exposed to mirrors, subjects showed socially appropriate behavior directed toward the mirror, as though the mirror image was another conspecific; and 2) as mirror exposure continued, the social behaviors waned and self directed/exploratory behaviors appeared. The appearance of self-directed behaviors was interpreted as indicative.

The second phase of this study will be conducted to further investigate the aforementioned behavior among a larger captive population of Asian elephants in California zoos and captive enclosures. In particular, this study will examine the presence or absence of vocalizations associated with self-recognition behavior. A detailed methodology of the study will be presented.
 
 

Professor Raman Sukumar, Centre for Ecological Sciences, Indian Institute of Science, Bangalore, India - Saturday, 4:00
The Asian Elephant: Conservation Biology of an Endangered Flagship Species

The Asian elephant, which once ranged from West Asia eastwards to northern Chine, is now confined to a 500,000 km2 area with a population of under 45,000 in the wild and 15,000 in captivity. Elephants are found in 13 Asian countries, with over 50% of the wild population in India. This charismatic megavertebrate which inhabits the biologically diverse forests of tropical Asia and has deep cultural links with the Asian peoples is possibly the ultimate flagship species for biodiversity conservation.

In this talk, I shall elaborate on three major issues that have the greatest relevance to the conservation of the Asian elephant. These are the loss and fragmentation of habitat, elephant human conflict at the forest-agriculture interface, and the poaching of elephants for ivory and other products.

The loss and fragmentation of the elephant's habitat is a historical process that still continues throughout its range. The result is the isolation of elephants in small populations with reduced viability in the face of demographic, genetic and environmental stochasticity. The preservation of corridors across elephant habitat landscapes should thus be an important consideration in conservation planning. Depredation of cultivated crops by elephants in small populations is widespread across Asia. The fragmentation of habitat is one proximate factor that increases the frequency of crop raiding. The ultimate cause of crop raiding seems to lie in the higher palatability and nutritive value of cultivated crops as compared to wild forage. The higher propensity of male elephants, as compared to female-led groups, to raid crops may be an outcome of a risk-taking strategy in males to enhance reproductive success that has been favored over evolutionary time, but may be maladaptive in this context. Strategies to minimize conflict, including barriers and population management, will be discussed.

It is not widely recognized that Asian elephant populations are also subjected to poaching for ivory and a variety of products including meat. Consequently, sex ratios are considerably skewed in favor of females in certain populations as in southern India. In other regions, such as Indo-China, entire populations have disappeared because of hunting for various products. The ecological consequences of poaching are discussed. With over 30% of the population in captivity, the proper care and management of captive stocks is obviously important for the overall conservation of the species.
 

Karen Willett, Department of Environmental Science and Policy, University of California, Davis - Saturday, 2:40
The Use of Geographic Information Systems (GIS) and Global Positioning Systems (GPS) for Elephant Monitoring and Population Estimates

Over the past decade years, the use of GIS and GPS technology for wildlife monitoring and population assessment has increased dramatically. The African Elephant Database (AED) was one of the earliest attempts to use GIS technology (Arc/Info) to estimate elephant populations for the African continent, and it is still in use today. A more recent application by Save the Elephants (STE) involves using GPS tracking collars to monitor elephant movements and analyze how they make decisions in relation to their needs. The Convention on International Trade of Endangered Species (CITES) is currently developing a monitoring system designed to measure and identify trends in the illegal killing of elephants in Africa and Asia, to determine changes in these trends and to assess whether and to what extent these trends are a result of changes in the status of African elephant populations within CITES. Several aspects of this program include use of GIS technology.
An overview of GIS and GPS technology, as well as a review of these and other applications of GIS and GPS for monitoring and estimating populations of elephants will be presented.
 

Lisa Wingate and Bill Lasley, University of California, Davis - Friday, 2:30
The Significance of Musth in Bull Elephants: Is It a Reproductive Event?

Musth is a complex behavioral and physiological phenomenon occurring in sexually mature bull elephants of both genera. It is known that musth involves a dramatic elevation in androgens. Musth bulls exhibit a number of behavioral and physical changes. While some of these changes may serve as signals for conspecifics, or increase the bull's likelihood of finding estrous females, they occur at considerable cost to the bull.

There are a number of advantages which bulls in musth enjoy. Musth bulls automatically attain a rank higher than any non-musth bull. Furthermore, there is evidence that musth bulls have more access to estrous females, and are in fact preferred by them as potential mates.

However, it is not clear that musth is purely a reproductive phenomenon. The evidence is unclear on whether or not the reproductive axis (hypothalamic-pituitary-gonadal) is responsible for the events of musth. Preliminary data from a pilot study will be presented which indicate increased adrenal activity during musth.

Alternative explanations for musth will be explored, and an experimental design detailed to address some of the questions raised.
 

George Wittemyer, Department of Environmental Science, Policy and Management, University of California, Berkeley - Saturday, 3:30
The Elephant Population of Samburu National Reserve, Kenya

A twenty-one month individual identification project on the Samburu and Buffalo Springs National Reserves' elephant population was conducted between November 1997 and August 1999. The population, of at least 749 elephants, was free ranging and relied heavily on areas outside the reserves. The number of elephants within the study area fluctuated daily. Reserves use was seasonal with the greatest numbers present during the wet season. Preliminary investigation of population social structure suggested that it consisted of two groups; resident and non-resident family units. The groups, comprised of approximately equal numbers of cows and calves, had different reserve use patterns and alternated breeding cycles. Census data uncovered a female biased adult population sex ratio, with nearly twice the number of females to males. The population was affected by poaching and political instability. Continued monitoring will assist conservation efforts by alerting authorities of major demographic or range use changes.