Elephant
Bibliographic
Database

.

...

Elephant Bibliographic Database
www.elephantcare.org

References updated October 2009 by date of publication, most recent first.

2009. Methicillin-resistant Staphylococcus aureus skin infections from an elephant calf--San Diego, California, 2008
91. MMWR Morb. Mortal. Wkly. Rep. 58, 194-198.
Abstract: Methicillin-resistant Staphylococcus aureus (MRSA) infections are a major cause of human skin and soft tissue infections in the United States. MRSA colonization and infection also have been observed in turtles, bats, seals, sheep, rabbits, rodents, cats, dogs, pigs, birds, horses, and cattle, and MRSA infections with an epidemiologic link to animal contact have been reported in veterinary personnel, pet owners, and farm animal workers. On January 29, 2008, the County of San Diego Health and Human Services Agency was notified of skin pustules on an African elephant (Loxodonta africana) calf and three of its caretakers at a zoo in San Diego County. After each of these infections (including the calf's infection) was laboratory confirmed as MRSA, an outbreak investigation and response was initiated by the zoo and the agency. This report summarizes the results of that investigation, which identified two additional confirmed MRSA infections, 15 suspected MRSA infections, and three MRSA-colonized persons (all among calf caretakers), and concluded that infection of the elephant calf likely came from a colonized caretaker. This is the first reported case of MRSA in an elephant and of suspected MRSA transmission from an animal to human caretakers at a zoo. Recommendations for preventing MRSA transmission in zoo settings include 1) training employees about their risks for infection and the recommended work practices to reduce them; 2) performing proper hand hygiene before and after animal contact; 3) using personal protective equipment (PPE) when working with ill or infected animals, especially during wound treatment; and 4) cleaning and disinfecting contaminated equipment and surfaces

Bates, L.A., Lee, P.C., Njiraini, N., Poole, J.H., Sayialel, K., Moss, C.J., Byrne, R., 2009. Do elephants show empathy? JOURNAL OF CONSCIOUSNESS STUDIES 15, 204-225.
Abstract: Elephants show a rich social organization and display a number of unusual traits. In this paper, we analyse reports collected over a thirty-five year period, describing behaviour that has the potential to reveal signs of empathic understanding. These include coalition formation, the offering of protection and comfort to others, retrieving and 'babysitting' calves, aiding individuals that would otherwise have difficulty in moving, and removing foreign objects attached to others. These records demonstrate that an elephant is capable of diagnosing animacy and goal directedness, and is able to understand the physical competence, emotional state and intentions of others, when they differ from its own. We argue that an empathic understanding of others is the simplest explanation of these abilities, and discuss reasons why elephants appear to show empathy more than other non-primate species.

Brown, J.L., Kersey, D.C., Freeman, E.W., Wagener, T., 2009. Assessment of diurnal urinary cortisol excretion in Asian and African elephants using different endocrine methods. Zoo. Biol.
Abstract: Longitudinal urine samples were collected from Asian and African elephants to assess sample processing and immunoassay techniques for monitoring adrenal activity. Temporal profiles of urinary cortisol measured by RIA and EIA, with and without dichloromethane extraction, were similar; all correlation coefficients were >0.90. However, based on regression analyses, cortisol immunoactivity in extracted samples was only 72-81% of that of unextracted values. Within assay technique, RIA values were only 74-81% of EIA values. Collection of 24-hr urine samples demonstrated a clear diurnal pattern of glucocorticoid excretion, with the lowest concentrations observed just before midnight and peak concentrations occurring around 0600-0800 hr. These results indicate that elephants fit the pattern of a diurnal species, and that glucocorticoid production is affected by a sleep-wake cycle similar to that described for other terrestrial mammals. Cortisol can be measured in both extracted and unextracted urine using RIA and EIA methodologies. However, unexplained differences in quantitative results suggest there may be sample matrix effects and that data generated using different techniques may not be directly comparable or interchangeable. Zoo Biol 28:1-10, 2009. (c) 2009 Wiley-Liss, Inc

Campos-Arceiz, A., Larrinaga, A.R., Weerasinghe, U.R., Takatsuki, S., Pastorini, J., Leimgruber, P., Fernand, P., Santamaria, L., 2009. Behavior rather than diet mediates seasonal differences in seed dispersal by Asian elephants. Ecology 89, 2684-2691.
Abstract: Digestive physiology and movement patterns of animal dispersers determine deposition patterns for endozoochorously dispersed seeds. We combined data from feeding trials, germination tests, and GPS telemetry of Asian elephants (Elephas maximus) to (1) describe the spatial scale at which Asian elephants disperse seeds; (2) assess whether seasonal differences in diet composition and ranging behavior translate into differences in seed shadows; and (3) evaluate whether scale and seasonal patterns vary between two ecologically distinct areas: Sri Lanka's dry monsoon forests and Myanmar's (Burma) mixed-deciduous forests. The combination of seed retention times (mean 39.5 h, maximum 114 h) and elephant displacement rates (average 1988 m in 116 hours) resulted in 50% of seeds dispersed over 1.2 km (mean 1222-2105 m, maximum 5772 m). Shifts in diet composition did not affect gut retention time and germination of ingested seeds. Elephant displacements were slightly longer, with stronger seasonal variation in Myanmar. As a consequence, seed dispersal curves varied seasonally with longer distances during the dry season in Myanmar but not in Sri Lanka. Seasonal and geographic variation in seed dispersal curves was the result of variation in elephant movement patterns, rather than the effect of diet changes on the fate of ingested seeds.

Cerling, T.E., Wittemyer, G., Ehleringer, J.R., Remien, C.H., Douglas-Hamilton, I., 2009. History of Animals using Isotope Records (HAIR): a 6-year dietary history of one family of African elephants
76. Proc. Natl. Acad. Sci. U. S. A 106, 8093-8100.
Abstract: The dietary and movement history of individual animals can be studied using stable isotope records in animal tissues, providing insight into long-term ecological dynamics and a species niche. We provide a 6-year history of elephant diet by examining tail hair collected from 4 elephants in the same social family unit in northern Kenya. Sequential measurements of carbon, nitrogen, and hydrogen isotope rations in hair provide a weekly record of diet and water resources. Carbon isotope ratios were well correlated with satellite-based measurements of the normalized difference vegetation index (NDVI) of the region occupied by the elephants as recorded by the global positioning system (GPS) movement record; the absolute amount of C(4) grass consumption is well correlated with the maximum value of NDVI during individual wet seasons. Changes in hydrogen isotope ratios coincided very closely in time with seasonal fluctuations in rainfall and NDVI whereas diet shifts to relatively high proportions of grass lagged seasonal increases in NDVI by approximately 2 weeks. The peak probability of conception in the population occurred approximately 3 weeks after peak grazing. Spatial and temporal patterns of resource use show that the only period of pure browsing by the focal elephants was located in an over-grazed, communally managed region outside the protected area. The ability to extract time-specific longitudinal records on animal diets, and therefore the ecological history of an organism and its environment, provides an avenue for understanding the impact of climate dynamics and land-use change on animal foraging behavior and habitat relations

Coeling, A., 2009. The application of social network theory to animal behavior. Bioscience Horizons 2, 32-43.
Abstract: Social network analysis (SNA) is a mathematical technique for analysing social relationships and the patterns and implications of these relationships (Wasserman S, Faust K (1994) Social Network Analysis: Methods and Applications. Cambridge: Cambridge University Press). It has only recently been discovered by behavioural biologists as a useful tool in the study of animal behaviour (Wey T, Blumstein DT, Shen W et al. (2008) Social network analysis of animal behaviour: a promising tool for the study of sociality. Anim Behav 75: 333-344). Video recording over a 2 month period was used to record the behaviour of the elephant group at Chester Zoo. SNA was applied in an investigation of the group structure and interactions of the group. Observations of individual and group behaviour were based upon 40 h of playback of the social interactions were recorded and analysed using AGNA (2003) and Pajek (2005) packages. The analysis showed that the many facets of individual behaviour could be understood in terms of social structure of the group. This study has demonstrated that SNA is a powerful approach to understanding group dynamics and is particularly applicable to the study of obligate social species. In conclusion, it is suggested that SNA is potentially a useful tool in the management of captive animal populations.

Freeman, E.W., Schulte, B.A., Brown, J.L., 2009. Using behavioral observations and keeper questionnaires to assess social relationships among captive female African elephants
60. Zoo. Biol. 28, 1-14.
Abstract: Free-ranging African elephants are highly social animals that live in a society where age, size, kinship, and disposition all contribute to social rank. Although captive elephant herds are small and largely comprises of unrelated females, dominance hierarchies are common. The goal of this study was to delineate how the behavior of captive female African elephants varies with respect to age and social rank based on a combination of keeper questionnaires and behavioral observations. "Body movements" and "trunk to" behaviors of 33 nonpregnant female African elephants housed at 14 North American zoos were recorded over 8 hr. Keepers at each facility also rated each elephant based on a series of questions about interactions with herdmates. The assessment of social rank based on observations correlated strongly with ranks assigned by keepers via the questionnaires. Observations and questionnaire responses indicated that body weight of the female, and to a lesser extent age, were significantly related to rates and types of "body movements" and that these demographic factors dictate the captive elephant hierarchy, similar to that observed in the wild. Many of the observed "body movements," such as back away, displace, push, and present, were correlated with keeper questionnaire responses about elephant interactions. However, none of the "trunk to" behaviors were related to age, size, or questionnaire responses even though they occurred frequently. In conclusion, we demonstrated that short-term behavioral observations and keeper questionnaires provided similar behavioral profiles for female African elephants housed in North American zoos. Zoo Biol 28:1-14, 2009. (c) 2009 Wiley-Liss, Inc

Freeman, E.W., Schulte, B.A., Brown, J.L., 2009. Investigating the impact of rank and ovarian activity on the social behavior of captive female African elephants . Zoo Biology 0, 1-14.
Abstract: Over a third of captive female African elephants in North America fail to exhibit normal estrous cycles based on long-term serum progestagen analyses. Why acyclicity occurs is unknown; however, the majority of noncycling females are ranked by keepers as the dominant individual within the group. To investigate the relationship between ovarian cyclicity status and keeper-determined social rank, observations were conducted on 33 female African elephants (18 cycling, 15 noncycling). Based on keeper evaluations, five cycling elephants were ranked dominant, seven in the middle and six as subordinate. In contrast, 10 noncycling elephants were ranked as dominate and five as subordinate with none ranked as middle. When comparing the behavior of the elephants by their keeper-determined rank, the dominant females dominant were significantly more likely to approach, displace and push. Similarly, keeper-determined subordinate females more frequently presented their hind end and held their ears erect. Behaviors initiated by one elephant toward another did not vary between cycling and noncycling females, except when the interaction with social rank was tested. Dominant, noncycling females initiated a higher percentage of approach and displace behaviors than both cycling and noncycling, subordinate elephants. Subordinate, noncycling elephants displayed the highest percentage of ears erect. Social rank drives the interactions of ex situ female African elephants more than ovarian cyclicity status. Thus, behavioral interactions cannot be used to predict which cycling elephants are most likely to become acyclic.

Freeman, E.W., Schulte, B.A., Brown, J.L., 2009. Investigating the impact of rank and ovarian activity on the social behavior of captive female African elephants . Zoo Biology 0, 1-14.
Abstract:
Over a third of captive female African elephants in North America fail to exhibit normal estrous cycles based on long-term serum progestagen analyses. Why acyclicity occurs is unknown; however, the majority of noncycling females are ranked by keepers as the dominant individual within the group. To investigate the relationship between ovarian cyclicity status and keeper-determined social rank, observations were conducted on 33 female African elephants (18 cycling, 15 noncycling). Based on keeper evaluations, five cycling elephants were ranked dominant, seven in the middle and six as subordinate. In contrast, 10 noncycling elephants were ranked as dominate and five as subordinate with none ranked as middle. When comparing the behavior of the elephants by their keeper-determined rank, the dominant females dominant were significantly more likely to approach, displace and push. Similarly, keeper-determined subordinate females more frequently presented their hind end and held their ears erect. Behaviors initiated by one elephant toward another did not vary between cycling and noncycling females, except when the interaction with social rank was tested. Dominant, noncycling females initiated a higher percentage of approach and displace behaviors than both cycling and noncycling, subordinate elephants. Subordinate, noncycling elephants displayed the highest percentage of ears erect. Social rank drives the interactions of ex situ female African elephants more than ovarian cyclicity status. Thus, behavioral interactions cannot be used to predict which cycling elephants are most likely to become acyclic.

Freeman, E.W., Guagnano, G., Olson, D., Keele, M., Brown, J.L., 2009. Social factors influence ovarian acyclicity in captive African elephants (Loxodonta africana). Zoo. Biol. 28, 1-15.
Abstract: Nearly one-third of reproductive age African elephants in North America that are hormonally monitored fail to exhibit estrous cycle activity, which exacerbates the nonsustainability of the captive population. Three surveys were distributed to facilities housing female African elephants to determine how social and environmental variables contribute to cyclicity problems. Forty-six facilities returned all three surveys providing information on 90% of the SSP population and 106 elephants (64 cycling, 27 noncycling and 15 undetermined). Logistic analyses found that some physiological and social history variables were related to ovarian acyclicity. Females more likely to be acyclic had a larger body mass index and had resided longer at a facility with the same herdmates. Results suggest that controlling the weight of an elephant might be a first step to helping mitigate estrous cycle problems. Data further show that transferring females among facilities has no major impact on ovarian activity. Last, social status appears to impact cyclicity status; at 19 of 21 facilities that housed both cycling and noncycling elephants, the dominant female was acyclic. Further studies on how social and environmental dynamics affect hormone levels in free-living, cycling elephants are needed to determine whether acyclicity is strictly a captivity-related phenomenon

Glaeser, S.G., Klinck, H., Mellinger, D.K., Ren, Y., 2009. A vocal repertoire of Asian elephants (Elephas maximus) and comparison of call classification methods. J Acoust Soc Am 125, 2710.
Abstract: This study compares classification methods applied to an acoustic repertoire of the Asian elephant (Elephas maximus). Recordings were made of captive elephants at the Oregon Zoo in Portland, OR 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 three methods. First, calls were classified manually using perceptual aural cues plus visual inspection of spectrograms for differentiation of fundamental frequency contour, tonality, and duration. Second, a set of 29 acoustic features was measured for nonoverlapping calls using the MATLAB-based program Osprey, then principal component analysis was applied to reduce the feature set. A neural network was used for classification. Finally, hidden Markov models, commonly used for pattern recognition, were utilized to recognize call types using perceptually-weighted cepstral features as input. All manual and automated classification methods agreed on structural distinction of six basic call types (trumpets, squeaks, squeals, roars, rumbles, and barks), with two call types (squeaks and squeals) being highly variable. Given the consistency of results among the classification methods across geographically and socially disparate subject groups, we believe automated call detection could successfully be applied to acoustic monitoring of Asian elephants.

Gobush, K., Kerr, B., Wasser, S., 2009. Genetic relatedness and disrupted social structure in a poached population of African elephants
110. Mol. Ecol. 18, 722-734.
Abstract: We use genetic measures of relatedness and observations of female bonding to examine the demographic signature of historically heavy poaching of a population of free-ranging African elephants. We collected dung samples to obtain DNA and observed behaviour from 102 elephant families over a 25-month period in 2003-2005 in Mikumi National Park, Tanzania. Poaching reduced the population by 75% in the decade prior to the 1989 ivory trade ban; park records indicate that poaching dropped significantly in Mikumi following the ban. Using 10 microsatellite loci, DNA was genotyped in 203 elephants and pair-wise relatedness was calculated among adult females within and between groups. The Mikumi population is characterized by small group size, considerable variation in group relatedness, females with no first-order adult relatives and females that form only weak social bonds. We used gene-drop analysis and a model of a genetically intact pedigree to compare our observed Mikumi group relatedness to a simulated genetically intact unpoached expectation. The majority of groups in Mikumi contain 2 to 3 adults; of these, 45% were classified as genetically disrupted. Bonding, quantified with a pair-wise association index, was significantly correlated with relatedness; however only half of the females formed strong bonds with other females, and relatedness was substantially lower for a given bond strength as compared to an unpoached population. Female African elephants without kin demonstrated considerable behavioural plasticity in this disturbed environment, grouping with other females lacking kin, with established groups, or remaining alone, unable to form any stable adult female-bonds. We interpret these findings as the remaining effect of poaching disturbance in Mikumi, despite a drop in the level of poaching since the commercial trade in ivory was banned 15 years ago

Hakeem, A.Y., Sherwood, C.C., Bonar, C.J., Butti, C., Hof, P.R., Allman, J.M., 2009. Von Economo neurons in the elephant brain. Anat. Rec. (Hoboken. ) 292, 242-248.
Abstract: Von Economo neurons (VENs), previously found in humans, all of the great ape species, and four cetacean species, are also present in African and Indian elephants. The VENs in the elephant are primarily found in similar locations to those in the other species. They are most abundant in the frontoinsular cortex (area FI) and are also present at lower density in the anterior cingulate cortex. Additionally, they are found in a dorsolateral prefrontal area and less abundantly in the region of the frontal pole. The VEN morphology appears to have arisen independently in hominids, cetaceans, and elephants, and may reflect a specialization for the rapid transmission of crucial social information in very large brains

Irie-Sugimoto, N., Kobayashi, T., Sato, T., Hasegawa, T., 2009. Relative quantity judgment by Asian elephants (Elephas maximus). Anim Cogn 12, 193-199.
Abstract: This study investigated whether Asian elephants can make relative quantity judgment (RQJ), a dichotomous judgment of unequal quantities ordered in magnitude. In Experiment 1, elephants were simultaneously shown two baskets with differing quantities of bait (up to 6 items). In Experiment 2, elephants were sequentially presented with baits, which could not be seen by elephants in their total quantities. The task of elephants was to choose the larger quantity in both experiments. Results showed that the elephants chose the larger quantity with significantly greater frequency. Interestingly, the elephants did not exhibit disparity or magnitude effects, in which performance declines with a smaller difference between quantities in a two-choice task, or the total quantity increases, respectively. These findings appear to be inconsistent with the previous reports of RQJ in other animals, suggesting that elephants may be using a different mechanism to compare and represent quantities than previously suggested for other species

Kun, A., Scheuring, I., 2009. Evolution of cooperation on dynamical graphs. Biosystems 96, 65-68.
Abstract:
There are two key characteristics of animal and human societies: (1) degree heterogeneity, meaning that not all individual have the same number of associates; and (2) the interaction topology is not static, i.e. either individuals interact with different set of individuals at different times of their life, or at least they have different associations than their parents. Earlier works have shown that population structure is one of the mechanisms promoting cooperation. However, most studies had assumed that the interaction network can be described by a regular graph (homogeneous degree distribution). Recently there are an increasing number of studies employing degree heterogeneous graphs to model interaction topology. But mostly the interaction topology was assumed to be static. Here we investigate the fixation probability of the cooperator strategy in the prisoner's dilemma, when interaction network is a random regular graph, a random graph or a scale-free graph and the interaction network is allowed to change.We show that the fixation probability of the cooperator strategy is lower when the interaction topology is described by a dynamical graph compared to a static graph. Even a limited network dynamics significantly decreases the fixation probability of cooperation, an effect that is mitigated stronger by degree heterogeneous networks topology than by a degree homogeneous one. We have also found that from the considered graph topologies the decrease of fixation probabilities due to graph dynamics is the lowest on scale-free graphs.

Leighty, K.A., Soltis, J., Wesolek, C.M., Savage, A., Mellen, J., Lehnhardt, J., 2009. GPS determination of walking rates in captive African elephants (Loxodonta africana)
79. Zoo. Biol. 28, 16-28.
Abstract: The movements of elephants in captivity have been an issue of concern for animal welfare activists and zoological professionals alike in recent years. In order to fully understand how movement rates reflect animal welfare, we must first determine the exact distances these animals move in the captive environment. We outfitted seven adult female African elephants (Loxodonta africana) at Disney's Animal Kingdom with collar-mounted global positioning recording systems to document their movement rates while housed in outdoor guest viewing habitats. Further, we conducted preliminary analyses to address potential factors impacting movement rates including body size, temperature, enclosure size, and social grouping complexity. We found that our elephants moved at an average rate of 0.409+/-0.007 km/hr during the 9-hr data collection periods. This rate translates to an average of 3.68 km traveled during the observation periods, at a rate comparable to that observed in the wild. Although movement rate did not have a significant relationship with an individual's body size in this herd, the movements of four females demonstrated a significant positive correlation with temperature. Further, females in our largest social group demonstrated a significant increase in movement rates when residing in larger enclosures. We also present preliminary evidence suggesting that increased social group complexity, including the presence of infants in the herd, may be associated with increased walking rates, whereas factors such as reproductive and social status may constrain movements

Mason, G.J., Veasey, J.S., 2009. How should the psychological well-being of zoo elephants be objectively investigated?
47. Zoo. Biol.
Abstract: Animal welfare (sometimes termed "well-being") is about feelings - states such as "suffering" or "contentment" that we can infer but cannot measure directly. Welfare indices have been developed from two main sources: studies of suffering humans, and of research animals deliberately subjected to challenges known to affect emotional state. We briefly review the resulting indices here, and discuss how well they are understood for elephants, since objective welfare assessment should play a central role in evidence-based elephant management. We cover behavioral and cognitive responses (approach/avoidance; intention, redirected and displacement activities; vigilance/startle; warning signals; cognitive biases, apathy and depression-like changes; stereotypic behavior); physiological responses (sympathetic responses; corticosteroid output - often assayed non-invasively via urine, feces or even hair; other aspects of HPA function, e.g. adrenal hypertrophy); and the potential negative effects of prolonged stress on reproduction (e.g. reduced gametogenesis; low libido; elevated still-birth rates; poor maternal care) and health (e.g. poor wound-healing; enhanced disease rates; shortened lifespans). The best validated, most used welfare indices for elephants are corticosteroid outputs and stereotypic behavior. Indices suggested as valid, partially validated, and/or validated but not yet applied within zoos include: measures of preference/avoidance; displacement movements; vocal/postural signals of affective (emotional) state; startle/vigilance; apathy; salivary and urinary epinephrine; female acyclity; infant mortality rates; skin/foot infections; cardio-vascular disease; and premature adult death. Potentially useful indices that have not yet attracted any validation work in elephants include: operant responding and place preference tests; intention and vacuum movements; fear/stress pheromone release; cognitive biases; heart rate, pupil dilation and blood pressure; corticosteroid assay from hair, especially tail-hairs (to access endocrine events up to a year ago); adrenal hypertrophy; male infertility; prolactinemia; and immunological changes. Zoo Biol 28:1-19, 2009. (c) 2009 Wiley-Liss, Inc

Pinter-Wollman, N., Isabell, L.A., Hart, L.A., 2009. Assessing translocation outcome: Comparing behavioral and physiological aspects of translocated and resident African elephants (Loxodonta africana). Biological Conservation 142, 1116-1124.
Abstract: Evaluating translocation outcomes is important for improving wildlife management and conservation actions. Often, when quick decisions need to be made and long-lived animals with slow reproductionrates are translocated, traditional assessment methods such as long-term survival and reproductive successcannot be used for assessing translocation outcomes. Thus, alternative, seldom used, measures suchas comparing the behavior and physiology of translocated animals to those of local residents should beemployed to assess the translocated animals' acclimation to their new home. Here we monitored the survival,physiology, and behavior of translocated African elephants (Loxodonta africana) and comparedthese measures to the local resident population at the release site. Adult male and female translocatedelephants' death rates were higher than those of the local population. Furthermore, the mortality rateof translocated adult males and calves was greater than expected based on their proportion in the translocatedelephant population. No difference was found in stress hormone levels between the two populations,but the body condition of the translocated elephants was significantly poorer than that of the localpopulation throughout the study period. The behavioral time budgets of the translocated elephants convergedwith those of the local population over time. Finally, translocated elephants utilized habitat thatwas similar to their source site (hills and permanent rivers) more than did the local population. Based on these findings we recommend careful consideration of timing, release location, and individuals targetedin future elephant translocations. More broadly, we introduce and explore seldom used translocation assessment techniques.

Pinter-Wollman, N., Isbell, L.A., Hart, L.A., 2009. The relationship between social behaviour and habitat familiarity in African elephants (Loxodonta africana)
117. Proc. Biol. Sci. 276, 1009-1014.
Abstract: Social associations with conspecifics can expedite animals' acclimation to novel environments. However, the benefits gained from sociality may change as the habitat becomes familiar. Furthermore, the particular individuals with whom animals associate upon arrival at a new place, familiar conspecifics or knowledgeable unfamiliar residents, may influence the type of information they acquire about their new home. To examine animals' social dynamics in novel habitats, we studied the social behaviour of African elephants (Loxodonta africana) translocated into a novel environment. We found that the translocated elephants' association with conspecifics decreased over time supporting our hypothesis that sociality provides added benefits in novel environments. In addition, we found a positive correlation between body condition and social association, suggesting that elephants gain direct benefits from sociality. Furthermore, the translocated elephants associated significantly less than expected with the local residents and more than expected with familiar, but not necessarily genetically related, translocated elephants. The social segregation between the translocated and resident elephants declined over time, suggesting that elephants can integrate into an existing social setting. Knowledge of the relationship between sociality and habitat familiarity is highly important in our constantly changing world to both conservation practice and our understanding of animals' behaviour in novel environments

Pinter-Wollman, N., 2009. Spatial behaviour of translocated African elephants (Loxodonta africana) in a novel environment: using behaviour to inform conservation actions. Behaviour 146, 1171-1192.
Abstract: When animals encounter a novel environment they can either reject it and leave or accept it and explore their new home. It is important to understand what governs animals' response to a novel place because of the fitness consequences and wildlife management implications
entailed. Here I examine the spatial behaviour of translocated African elephants (Loxodonta africana) upon arrival at a novel environment. I monitored the movement patterns of 12 radiocollared elephants for a year post-translocation. I documented the first account of both female
and male African elephants homing back to their natal habitat. More males than expected left the release site, but female-calf units also homed to their natal habitat, demonstrating that homing is not confined to one sex or age.When examining the spatial behaviour of elephants
that remained near the release site I did not find a relationship between habitat exploration and last distance from release site, elephant age, or social association. However, I did find a negative correlation between habitat exploration and distance from human activities. This
work provides biological insights regarding individual variation in spatial activity of animals in a novel environment and offers recommendations for future management actions.

Plotnik, J.M., de Waal, F.B., Moore, D., III, Reiss, D., 2009. Self-recognition in the Asian elephant and future directions for cognitive research with elephants in zoological settings
48. Zoo. Biol.
Abstract: The field of animal cognition has grown steadily for nearly four decades, but the primary focus has centered on easily kept lab animals of varying cognitive capacity, including rodents, birds and primates. Elephants (animals not easily kept in a laboratory) are generally thought of as highly social, cooperative, intelligent animals, yet few studies-with the exception of long-term behavioral field studies-have been conducted to directly support this assumption. In fact, there has been remarkably little cognitive research conducted on Asian (Elephas maximus) or African (Loxodonta africana or L. cyclotis) elephants. Here, we discuss the opportunity and rationale for conducting such research on elephants in zoological facilities, and review some of the recent developments in the field of elephant cognition, including our recent study on mirror self-recognition in E. maximus. Zoo Biol 28:1-13, 2009. (c) 2009 Wiley-Liss, Inc

Rees, P.A., 2009. Activity budgets and the relationship between feeding and stereotypic behaviors in Asian elephants (Elephas maximus) in a Zoo
75. Zoo. Biol. 28, 79-97.
Abstract: Activity budgets were studied in eight Asian elephants (Elephas maximus) at Chester Zoo (UK) for 35 days, between January and November 1999. Recordings were made between 10:00 and 16:00 hr (with most behavior frequencies calculated between 10:00 and 14:00 hr). The elephants exhibited variation in activity depending on their age, sex, the time of day and the time of year. Only the five adult cows exhibited stereotypic behavior, with frequencies ranging from 3.9 to 29.4% of all observations. These elephants exhibited individual, diurnal and seasonal variation in stereotypic behavior. This has implications for studies that use short sampling periods and may make comparisons of data collected at different times of the day or year invalid. The six adult elephants spent 27.4-41.4% of the time feeding (between 10:00 and 14:00 hr), 22.9-42.0% standing still, 6.1-19.2% walking and 3.9-9.6% dusting. The hypothesis that the frequency of stereotypic behavior in adult cow elephants was negatively correlated with the frequency of feeding behavior was tested and was found to be true. Stereotypic behavior increased in frequency toward the end of the day-while waiting to return to the elephant house for food--and elephants spent more time stereotyping during the winter months than during the summer months. Elephants were inactive (i.e. exhibited behaviors other than locomotion) for between 70.1 and 93.9% of the time. Creating more opportunities for elephants to exhibit foraging behavior and the introduction of greater unpredictability into management regimes, especially feeding times, may reduce the frequency of stereotypic behavior and increase general activity levels

Rees, P.A., 2009. The sizes of elephant groups in zoos: implications for elephant welfare. J. Appl. Anim Welf. Sci. 12, 44-60.
Abstract: This study examined the distribution of 495 Asian elephants (Elephas maximus) and 336 African elephants (Loxodonta africana) in 194 zoos, most of which were located in Europe (49.1%) and North America (32.6%). Cows outnumbered bulls 4 to 1 (Loxodonta) and 3 to 1 (Elephas). Groups contained 7 or fewer: mean, 4.28 (sigma = 5.73). One fifth of elephants lived alone or with one conspecific. Forty-six elephants (5.5%) had no conspecific. Many zoos ignore minimum group sizes of regional zoo association guidelines. The American Zoo and Aquarium Association recommends that breeding facilities keep herds of 6 to 12 elephants. The British and Irish Association of Zoos and Aquariums recommends keeping together at least 4 cows over 2 years old. Over 69% Asian and 80% African cow groups-including those under 2 years-consisted of fewer than 4 individuals. Recently, Europe and North America have made progress with some zoos no longer keeping elephants and with others investing in improved facilities and forming larger herds. The welfare of individual elephants should outweigh all other considerations; zoos should urgently seek to integrate small groups into larger herds

Remis, M.J., Hardin, R., 2009. Transvalued Species in an African Forest. Conserv. Biol.
Abstract: We combined ethnographic investigations with repeated ecological transect surveys in the Dzanga-Sangha Dense Forest Reserve (RDS), Central African Republic, to elucidate consequences of intensifying mixed use of forests. We devised a framework for transvaluation of wildlife species, which means the valuing of species on the basis of their ecological, economic, and symbolic roles in human lives. We measured responses to hunting, tourism, and conservation of two transvalued species in RDS: elephants (Loxodonta cyclotis) and gorillas (Gorilla gorilla). Our methods included collecting data on encounter rates and habitat use on line transects. We recorded cross-cultural variation in ideas about and interactions with these species during participant observation of hunting and tourism encounters and ethnographic interviews with hunters, conservation staff, researchers, and tourists. Ecologically, gorillas used human-modified landscapes successfully, and elephants were more vulnerable than gorillas to hunting. Economically, tourism and encounters with elephants and gorillas generated revenues and other benefits for local participants. Symbolically, transvaluation of species seemed to undergird competing institutions of forest management that could prove unsustainable. Nevertheless, transvaluation may also offer alternatives to existing social hierarchies, thereby integrating local and transnational support for conservation measures. The study of transvaluation requires attention to transnational flows of ideas and resources because they influence transspecies interactions. Cross-disciplinary in nature, transvalution of species addresses the political and economic challenges to conservation because it recognizes the varied human communities that shape the survival of wildlife in a given site. Transvaluation of species could foster more socially inclusive management and monitoring approaches attuned to competing economic demands, specific species behaviors, and human practices at local scales

Soltis, J., 2009. Vocal communication in African Elephants (Loxodonta africana)
61. Zoo. Biol. 28, 1-18.
Abstract: Research on vocal communication in African elephants has increased in recent years, both in the wild and in captivity, providing an opportunity to present a comprehensive review of research related to their vocal behavior. Current data indicate that the vocal repertoire consists of perhaps nine acoustically distinct call types, "rumbles" being the most common and acoustically variable. Large vocal production anatomy is responsible for the low-frequency nature of rumbles, with fundamental frequencies in the infrasonic range. Additionally, resonant frequencies of rumbles implicate the trunk in addition to the oral cavity in shaping the acoustic structure of rumbles. Long-distance communication is thought possible because low-frequency sounds propagate more faithfully than high-frequency sounds, and elephants respond to rumbles at distances of up to 2.5 km. Elephant ear anatomy appears designed for detecting low frequencies, and experiments demonstrate that elephants can detect infrasonic tones and discriminate small frequency differences. Two vocal communication functions in the African elephant now have reasonable empirical support. First, closely bonded but spatially separated females engage in rumble exchanges, or "contact calls," that function to coordinate movement or reunite animals. Second, both males and females produce "mate attraction" rumbles that may advertise reproductive states to the opposite sex. Additionally, there is evidence that the structural variation in rumbles reflects the individual identity, reproductive state, and emotional state of callers. Growth in knowledge about the communication system of the African elephant has occurred from a rich combination of research on wild elephants in national parks and captive elephants in zoological parks. Zoo Biol 28:1-18, 2009. (c) 2009 Wiley-Liss, Inc

Soltis, J., Leighty, K.A., Wesolek, C.M., Savage, A., 2009. The expression of affect in African elephant (Loxodonta africana) rumble vocalizations
59. J. Comp Psychol. 123, 222-225.
Abstract: Affective states are thought to be expressed in the mammalian voice, but such investigations are most common in primates. Source and filter features of rumbles were analyzed from 6 adult female African elephants (Loxodonta africana) at Disney's Animal Kingdom. Rumbles produced during periods of minimal social interaction ("low affect") were compared to those produced during dominance interactions ("high affect"). Low-ranking females produced rumbles with increased and more variable fundamental frequencies, and increased durations and amplitudes during dominance interactions with superiors, compared to the low affect context. This acoustic response is consistent with the expression of affect in mammals and may signal submission to superiors. The 2 highest ranking females were codominant and competed for alpha status. They produced rumbles with decreased and less variable fundamental frequencies, increased durations and amplitudes, and a decrease in formant dispersion during dominance interactions with each other, compared to the low affect context. This response is not generally consistent with the expression of affect, but may signal large body size to competitors. These results suggest that affect can be expressed in the voiced sounds of elephants

Thompson, M.E., Schwager, S.J., Payne, K.B., Turkalo, A.K., 2009. Acoustic estimation of wildlife abundance: methodology for vocal mammals in forested habitats. African Journal of Ecology.
Abstract: Habitat loss and hunting pressure threaten mammal populations worldwide, generating critical time constraints on trend assessment. This study introduces a new survey method that samples continuously and non-invasively over long time periods, obtaining estimates of abundance from vocalization rates. We present feasibility assessment methods for acoustic surveys and develop equations for estimating population size. As an illustration, we demonstrate the feasibility of acoustic surveys for African forest elephants (Loxodonta africana cyclotis). Visual surveys and vocalizations from a forest clearing in the Central African Republic were used to establish that low-frequency elephant calling rate is a useful index of elephant numbers (linear regressionP<0.001,radj.2=0.58). The effective sampling area was 3.22km2per acoustic sensor, a dramatic increase in coverage over dung survey transects. These results support the use of acoustic surveys for estimating elephant abundance over large remote areas and in diverse habitats, using a distributed network of acoustic sensors. The abundance estimation methods presented can be applied in surveys of any species for which an acoustic abundance index and detection function have been established. This acoustic survey technique provides an opportunity to improve management and conservation of many acoustically-active taxa whose populations are currently under-monitored.

Wesolek, C.M., Soltis, J., Leighty, K.A., Savage, A., 2009. Infant African elephan rumble vocalizations vary according to social interactions with adult females. Bioacoustics 18, 227-239.
Abstract: Research on African elephant (Loxodonta africana) vocal communication has increased in recent years, yet there has been very little data collected on the vocal production of infant African elephants. Vocalizations were recorded from a group of five adult female African elephants and 3 dependent offspring (1 male and 2 female) at Disney's Animal Kingdom, Florida, U.S.A., using custom-designed audio-recording collars worn by the adult females. We measured both source and filter features of infant 'rumble' vocalizations made during affiliative social interactions and after cessation of nursing from adult females. Rumble vocalizations produced in the 'nurse cessation' context exhibited an upward shift in formant frequency locations, compared to rumbles produced during the 'affiliation' context. Additionally, call duration increased and fundamental frequencies decreased after nurse cessations for the male, but both females showed the opposite acoustic response. When infant rumbles accompanied nurse cessations, nursing was more likely to resume within 30 seconds compared to nurse cessations without vocalizations. These results suggest that infant rumbles associated with cessation of nursing reflect the motivational state of infants and may influence maternal responsiveness.

Wittemyer, G., Okello, J.B., Rasmussen, H.B., Arctander, P., Nyakaana, S., Douglas-Hamilton, I., Siegismund, H.R., 2009. Where sociality and relatedness diverge: the genetic basis for hierarchical social organization in African elephants. Proc Royal Soc Biol 276, 3513-3521.
Abstract: Hierarchical properties characterize elephant fission-fusion social organization whereby stable groups of individuals coalesce into higher order groups or split in a predictable manner. This hierarchical complexity is rare among animals and, as such, an examination of the factors driving its emergence offers unique insight into the evolution of social behaviour. Investigation of the genetic basis for such social affiliation demonstrates that while the majority of core social groups (second-tier affiliates) are significantly related, this is not exclusively the case. As such, direct benefits received through membership of these groups appear to be salient to their formation and maintenance. Further analysis revealed that the majority of groups in the two higher social echelons (third and fourth tiers) are typically not significantly related. The majority of third-tier members are matrilocal, carrying the same mtDNA control region haplotype, while matrilocality among fourth-tier groups was slightly less than expected at random. Comparison of results to those from a less disturbed population suggests that human depredation, leading to social disruption, altered the genetic underpinning of social relations in the study population. These results suggest that inclusive fitness benefits may crystallize elephant hierarchical social structuring along genetic lines when populations are undisturbed. However, indirect benefits are not critical to the formation and maintenance of second-, third- or fourth-tier level bonds, indicating the importance of direct benefits in the emergence of complex, hierarchical social relations among elephants. Future directions and conservation implications are discussed

Bates, L.A., Poole, J.H., Byrne, R.W., 2008. Elephant cognition. Curr. Biol. 18, R544-R546.

Evans, K., Harris, S., 2008. Adolescence in male African elephants, Loxodonta africana, and the importance of sociality. Animal Behavior 76, 779-787.
Abstract: The degree of sociality during an animal's life changes as it modulates its behaviour to reflect different lifestages. Only a few species of mammal undergo a period of adolescence, but for these species it is probablyone of their most important life stages. It is when individuals acquire skills and develop relationships thatare of both immediate and long-term benefit to their survival and reproductive success, particularly in polygynous males in which sexual selection favours size and dominance. We collected focal and observationaldata on male African elephants in the Okavango Delta, Botswana, to assess behaviour and socialinteractions during adolescence. Adolescent males (10-15 and 16-20 years of age) were the most sociableage group, showing preferences for larger social groupings and being in closer proximity to other elephants;later adolescent males (ages 16-20) showed a tendency for higher social levels. Males of all agespreferred to have males 36 years of age as their nearest neighbour. We argue that this proximity to oldermales provides opportunities for males to learn from more experienced individuals. It has long been recognized that matriarchs are the repositories of social and ecological knowledge within elephant breedingherds: we suggest that mature males are reservoirs for such knowledge within bull society.

Gobush, K.S., Mutayoba, B.M., Wasser, S.K., 2008. Long-term impacts of poaching on relatedness, stress physiology, and reproductive output of adult female african elephants. Conserv. Biol. 22, 1590-1599.
Abstract: Widespread poaching prior to the 1989 ivory ban greatly altered the demographic structure of matrilineal African elephant (Loxodonta africana) family groups in many populations by decreasing the number of old, adult females. We assessed the long-term impacts of poaching by investigating genetic, physiological, and reproductive correlates of a disturbed social structure resulting from heavy poaching of an African elephant population in Mikumi National Park, Tanzania, prior to 1989. We examined fecal glucocorticoid levels and reproductive output among 218 adult female elephants from 109 groups differing in size, age structure, and average genetic relatedness over 25 months from 2003 to 2005. The distribution in group size has changed little since 1989, but the number of families with tusked old matriarchs has increased by 14.2%. Females from groups that lacked an old matriarch, first-order adult relatives, and strong social bonds had significantly higher fecal glucocorticoid values than those from groups with these features (all females R(2)= 0.31; females in multiadult groups R(2)= 0.46). Females that frequented isolated areas with historically high poaching risk had higher fecal glucocorticoid values than those in low poaching risk areas. Females with weak bonds and low group relatedness had significantly lower reproductive output (R(2)[U]=0.21). Females from disrupted groups, defined as having observed average group relatedness 1 SD below the expected mean for a simulated unpoached family, had significantly lower reproductive output than females from intact groups, despite many being in their reproductive prime. These results suggest that long-term negative impacts from poaching of old, related matriarchs have persisted among adult female elephants 1.5 decades after the 1989 ivory ban was implemented

Hill, R.A., Bentley, R.A., Dunbar, R.I., 2008. Network scaling reveals consistent fractal pattern in hierarchical mammalian societies. Biol. Lett. 4, 748-751.
Abstract: Recent studies have demonstrated that human societies are hierarchically structured with a consistent scaling ratio across successive layers of the social network; each layer of the network is between three and four times the size of the preceding (smaller) grouping level. Here we show that similar relationships hold for four mammalian taxa living in multi-level social systems. For elephant (Loxodonta africana), gelada (Theropithecus gelada) and hamadryas (Papio hamadryas hamadryas) baboon, successive layers of social organization have a scaling ratio of almost exactly 3, indicating that such branching ratios may be a consistent feature of all hierarchically structured societies. Interestingly, the scaling ratio for orca (Orcinus orca) was 3.8, which might mean that aquatic environments place different constraints on the organization of social hierarchies. However, circumstantial evidence from a range of other species suggests that scaling ratios close to 3 may apply widely, even in species where hierarchical social structures have not traditionally been identified. These results identify the origin of the hierarchical, fractal-like organization of mammalian social systems as a fundamental question

Meyer, J., Goodwin, T., Schulte, B., 2008. Intrasexual chemical communication and social responses ofcaptive female African elephants. Animal Behavior 76, 163-174.
Abstract: In matrilineal societies, competition between females can occur within and between social units. Dominance hierarchies reduce costly conflicts when reliable cues of status are available, and reproductive condition may alter individual or group status. Female African elephants live in matriarchal groups with linear dominance hierarchies occurring within and between groups; elephants use chemical signals to mediate social interactions. If reproductive condition has important implications for inter- or intragroup behaviour, then females should discriminate between chemical signal sources that reveal reproductive condition. We examined whether trunk-tip contacts between females within a social group were related to phase of oestrus. Observations were conducted on 21 reproductively viable females at nine zoological facilities in North America. Females in the follicular phase received contacts to the urogenital region at a higher rate than did luteal phase females, and contacts increased with approaching ovulation. This supports the existence of an oestrous signal. We also examined whether an oestrous signal was evident by female investigation of urine collected from the luteal and follicular phases of unfamiliar conspecifics. Elephants responded to unfamiliar urine more than to the control, but response rates to the urine types did not differ. Females within a social unit detected differences in oestrus, but they did not show such discrimination to urinary signals from unfamiliar females. Further evaluation of the existence of a female-to-female oestrous pheromone requires assessing responses to urine from familiar individuals. Understanding the relationship between oestrous condition and dominance status can shed light on the adaptive value of sociality.

Okello, J.B., Masembe, C., Rasmussen, H.B., Wittemyer, G., Omondi, P., Kahindi, O., Muwanika, V.B., Arctander, P., Douglas-Hamilton, I., Nyakaana, S., Siegismund, H.R., 2008. Population genetic structure of savannah elephants in Kenya: conservation and management implications. J. Hered. 99, 443-452.
Abstract: We investigated population genetic structure and regional differentiation among African savannah elephants in Kenya using mitochondrial and microsatellite markers. We observed mitochondrial DNA (mtDNA) nucleotide diversity of 1.68% and microsatellite variation in terms of average number of alleles, expected and observed heterozygosities in the total study population of 10.20, 0.75, and 0.69, respectively. Hierarchical analysis of molecular variance of mtDNA variation revealed significant differentiation among the 3 geographical regions studied (F(CT) = 0.264; P < 0.05) and a relatively lower differentiation among populations within regions (F(SC) = 0.218; P < 0.0001). Microsatellite variation significantly differentiated among populations within regions (F(SC) = 0.019; P < 0.0001) but not at the regional levels (F(CT) = 0.000; P > 0.500). We attribute the high differentiation at the mitochondrial genome to the matrilineal social structure of elephant populations, female natal philopatry, and probably ancient vicariance. Lack of significant regional differentiation at the nuclear loci vis-a-vis strong differences at mtDNA loci between regions is likely the effect of subsequent homogenization through male-mediated gene flow. Our results depicting 3 broad regional mtDNA groups and the observed population genetic differentiation as well as connectivity patterns should be incorporated in the planning of future management activities such as translocations

Rasmssen, H.B., Ganswindt, A., Douglas-Hamilton, I., Vollrath, F., 2008. Endocrine and behavioral changes in male African elephants: Linking hormone changes to sexual state and reproductive tactics. Hormones and Behavior.
Abstract: Endocrine and behavioral changes in male African elephants: Linking hormone changes to sexual state and reproductive tactics.Henrik B Rasmussen, Andre Ganswindt, Iain Douglas-Hamilton, and Fritz VollrathHormones and Behavior, May 22, 2008                       
Hormones play a crucial role in mediating genetic and environmental effects into morphological and behavioral phenotypes. In systems with alternative reproductive tactics (ART) shifts between tactics are hypothesized to be under proximate hormonal control. Most studies of the underlying endocrine changes behind ART have focused on fish and amphibians rather than mammals and few have investigated the potential interaction between different endocrine axes in regulating shifts between conditional dependent tactics. Using a combination of endocrine and behavioral data from male African elephants we expand on our previously published analysis and show that the initial increase in androgens predates the behavioral shifts associated with reproductively active periods, supporting the role of androgens in activating sexually active periods in males. A strong interactive effect between androgens and glucocorticoids was found to determine the presence or absence of temporal gland secretion and urine dribbling, signals associated with the competitive reproductive tactic of musth, with elevated glucocorticoids levels suppressing the occurrence of musth signals. In addition external environmental conditions affected hormone levels. The presence of receptive females resulted in elevated androgens in dominant musth males but increased glucocorticoids in subordinate non-musth males. The presented data on hormones, behavior and reproductive tactics strongly support an underlying endocrine mechanism for mediating the translation of intrinsic as well as extrinsic local conditions into the conditional dependent reproductive tactics in male elephants via interactions between the hypothalamic-pituitary-gonadal and -adrenal axes.

Rasmussen, H.B., Ganswindt, A., Douglas-Hamilton, I., Vollrath, F., 2008. Endocrine and behavioral changes in male African elephants: linking hormone changes to sexual state and reproductive tactics. Horm. Behav. 54, 539-548.
Abstract: Hormones play a crucial role in mediating genetic and environmental effects into morphological and behavioral phenotypes. In systems with alternative reproductive tactics (ART) shifts between tactics are hypothesized to be under proximate hormonal control. Most studies of the underlying endocrine changes behind ART have focused on fish and amphibians rather than mammals and few have investigated the potential interaction between different endocrine axes in regulating shifts between conditional dependent tactics. Using a combination of endocrine and behavioral data from male African elephants we expand on our previously published analysis and show that the initial increase in androgens predates the behavioral shifts associated with reproductively active periods, supporting the role of androgens in activating sexually active periods in males. A strong interactive effect between androgens and glucocorticoids was found to determine the presence or absence of temporal gland secretion and urine dribbling, signals associated with the competitive reproductive tactic of musth, with elevated glucocorticoids levels suppressing the occurrence of musth signals. In addition external environmental conditions affected hormone levels. The presence of receptive females resulted in elevated androgens in dominant musth males but increased glucocorticoids in subordinate non-musth males. The presented data on hormones, behavior and reproductive tactics strongly support an underlying endocrine mechanism for mediating the translation of intrinsic as well as extrinsic local conditions into the conditional dependent reproductive tactics in male elephants via interactions between the hypothalamic-pituitary-gonadal and -adrenal axes

Shannon, G., Page, B.R., Mackey, R.L., Duffy, K.J., Slotow, R., 2008. Activity budgets and sexual segregation in African elephants (Loxodonta africana). Journal of Mammalogy 89, 467-476.
Abstract: The activity budget hypothesis is 1 of 4 main hypotheses proposed to explain sexual segregation by large herbivores. Because of their smaller body size, females are predicted to have higher mass-specific energy requirements and lower digestive efficiency than males. As a result, females are expected to forage longer to satisfy their nutritional demands. Maintaining the cohesion of a mixed-sex group with differing activity budgets and asynchronous behavioral patterns is increasingly difficult, ultimately leading to spatial segregation of males and females. We tested this hypothesis using data (2002-2005) from 3 distinct populations of African elephants (Loxodonta africana), a species that exhibits marked sexual segregation. Group and individual behaviors were assessed at discrete points in time throughout the day, with a minimum of 10 min between consecutive records. Focal samples of individual male and female elephants also were recorded, with behavioral data logged every minute for 15 min. Data were grouped into 5 behavioral categories: drinking, resting, walking, feeding, and other. Neither activity rhythms nor feeding time varied significantly between the sexes and behavioral patterns were very similar. We propose that social and environmental factors influence behavioral rhythms to a greater extent than does body size, whereas increasing feeding time is only 1 method by which elephants can improve nutritional return. This is especially pertinent when considering their generalist foraging approach, substantial energy demands, and hindgut fermentation. We conclude that the activity budget hypothesis is unlikely to be the causal mechanism in the sexual segregation of African elephants, a finding that concurs with recent experimental and field research on a range of sexually dimorphic herbivores.

Slade-Cain, B.E., Rasmussen, L.E., Schulte, B.A., 2008. Estrous state influences on investigative, aggressive, and tail flicking behavior in captive female Asian elephants
78. Zoo. Biol. 27, 167-180.
Abstract: Females of species that live in matrilineal hierarchies may compete for temporally limited resources, yet maintain social harmony to facilitate cohesion. The relative degree of aggressive and nonaggressive interactions may depend on the reproductive condition of sender and receiver. Individuals can benefit by clearly signaling and detecting reproductive condition. Asian elephants (Elephas maximus) live in social matrilineal herds. Females have long estrous cycles (14-16 weeks) composed of luteal (8-12 weeks) and follicular (4-8 weeks) phases. In this study, we observed the behavior of four captive Asian elephant females during multiple estrous cycles over 2 years. We evaluated whether investigative, aggressive, and tail flicking behaviors were related to reproductive condition. Investigative trunk tip contacts showed no distinct pattern by senders, but were more prevalent toward female elephants that were in their follicular compared with their luteal phase. The genital area was the most frequently contacted region and may release reproductively related chemosignals. Aggression did not differ significantly with estrus; however, rates of aggression were elevated when senders were approaching ovulation and receivers were in the luteal phase. Females in the follicular phase may honestly advertise their condition. Contacts by conspecifics may serve to assess condition and reduce aggression. A behavior termed "tail flicking" was performed mainly during the mid-follicular phase when estrogen and luteinizing hormone levels are known to spike. Tail flicking may disperse chemical signals in urine or mucus as well as act as a tonic signal that could provide a means of anticipating forthcoming ovulation by elephants and also for human observers and caretakers. Zoo Biol 27:167-180, 2008. (c) 2008 Wiley-Liss, Inc

Tyack, P.L., 2008. Convergence of calls as animals form social bonds, active compensation for noisy communication channels, and the evolution of vocal learning in mammals. J. Comp Psychol. 122, 319-331.
Abstract: The classic evidence for vocal production learning involves imitation of novel, often anthropogenic sounds. Among mammals, this has been reported for dolphins, elephants, harbor seals, and humans. A broader taxonomic distribution has been reported for vocal convergence, where the acoustic properties of calls from different individuals converge when they are housed together in captivity or form social bonds in the wild. Vocal convergence has been demonstrated for animals as diverse as songbirds, parakeets, hummingbirds, bats, elephants, cetaceans, and primates. For most species, call convergence is thought to reflect a group-distinctive identifier, with shared calls reflecting and strengthening social bonds. A ubiquitous function for vocal production learning that is starting to receive attention involves modifying signals to improve communication in a noisy channel. Pooling data on vocal imitation, vocal convergence, and compensation for noise suggests a wider taxonomic distribution of vocal production learning among mammals than has been generally appreciated. The wide taxonomic distribution of this evidence for vocal production learning suggests that perhaps more of the neural underpinnings for vocal production learning are in place in mammals than is usually recognized

Wittemyer, G., Polansky, L., Douglas-Hamilton, I., Getz, W.M., 2008. Disentangling the effects of forage, social rank, and risk on movement autocorrelation of elephants using Fourier and wavelet analyses. Proc. Natl. Acad. Sci. U. S. A 105, 19108-19113.
Abstract: The internal state of an individual-as it relates to thirst, hunger, fear, or reproductive drive-can be inferred by referencing points on its movement path to external environmental and sociological variables. Using time-series approaches to characterize autocorrelative properties of step-length movements collated every 3 h for seven free-ranging African elephants, we examined the influence of social rank, predation risk, and seasonal variation in resource abundance on periodic properties of movement. The frequency domain methods of Fourier and wavelet analyses provide compact summaries of temporal autocorrelation and show both strong diurnal and seasonal based periodicities in the step-length time series. This autocorrelation is weaker during the wet season, indicating random movements are more common when ecological conditions are good. Periodograms of socially dominant individuals are consistent across seasons, whereas subordinate individuals show distinct differences diverging from that of dominants during the dry season. We link temporally localized statistical properties of movement to landscape features and find that diurnal movement correlation is more common within protected wildlife areas, and multiday movement correlations found among lower ranked individuals are typically outside of protected areas where predation risks are greatest. A frequency-related spatial analysis of movement-step lengths reveal that rest cycles related to the spatial distribution of critical resources (i.e., forage and water) are responsible for creating the observed patterns. Our approach generates unique information regarding the spatial-temporal interplay between environmental and individual characteristics, providing an original approach for understanding the movement ecology of individual animals and the spatial organization of animal populations

Bates, L.A., Sayialel, K.N., Nijiraini, N.W., Moss, C.J., Poole, J.H., Byrne, R.W., 2007. Elephants classify human ethnic groups by odor and garment color. Current Biology 17, 1-5.
Abstract: PrFont34Bin0BinSub0Frac0Def1Margin0Margin0Jc1Indent1440Lim0Lim1Animals can benefit from classifying predators or other dangers into categories, tailoring their escape strategies to the type and nature of the risk. Studies of alarm vocalizations have revealed various levels of sophistication in classification [1-5]. In many taxa, reactions to danger are inflexible, but some species can learn the level of threat presented by the local population of a predator [6-8] or by specific, recognizable individuals [9, 10]. Some species distinguish several species of predator, giving differentiated warning calls and escape reactions; here, we explore an animal's classification of subgroups within a species. We show that elephants distinguish at least two Kenyan ethnic groups and can identify them by olfactory and color cues independently. In the Amboseli ecosystem, Kenya, young Maasai men demonstrate virility by spearing elephants (Loxodonta africana), but Kamba agriculturalists pose little threat. Elephants showed greater fear when they detected the scent of garments previously worn by Maasai than by Kamba men, and they reacted aggressively to the color associated with Maasai. Elephants are therefore able to classify members of a single species into subgroups that pose different degrees of danger.

Bates, L.A., Byrne, R.W., 2007. Creative or created: using anecdotes to investigate animal cognition. Methods 42, 12-21.
Abstract: In non-human animals, creative behaviour occurs spontaneously only at low frequencies, so is typically missed by standardised observational methods. Experimental approaches have tended to rely overly on paradigms from child development or adult human cognition, which may be inappropriate for species that inhabit very different perceptual worlds and possess quite different motor capacities than humans. The analysis of anecdotes offers a solution to this impasse, provided certain conditions are met. To be reliable, anecdotes must be recorded immediately after observation, and only the records of scientists experienced with the species and the individuals concerned should be used. Even then, interpretation of a single record is always ambiguous, and analysis is feasible only when collation of multiple records shows that a behaviour pattern occurs repeatedly under similar circumstances. This approach has been used successfully to study a number of creative capacities of animals: the distribution, nature and neural correlates of deception across the primate order; the occurrence of teaching in animals; and the neural correlates of several aptitudes--in birds, foraging innovation, and in primates, innovation, social learning and tool-use. Drawing on these approaches, we describe the use of this method to investigate a new problem, the cognition of the African elephant, a species whose sheer size and evolutionary distance from humans renders the conventional methods of comparative psychology of little use. The aim is both to chart the creative cognitive capacities of this species, and to devise appropriate experimental methods to confirm and extend previous findings

Bates, L.W., Byrne, R.W., 2007. Creative or created: Using anecdotes to investigate animal cognition. Methods 42, 12-21.
Abstract: In non-human animals, creative behaviour occurs spontaneously only at low frequencies, so is typically missed by standardised observational methods. Experimental approaches have tended to rely overly on paradigms from child development or adult human cognition, which may be inappropriate for species that inhabit very different perceptual worlds and possess quite different motor capacities than humans. The analysis of anecdotes offers a solution to this impasse, provided certain conditions are met. To be reliable, anecdotes must be recorded immediately after observation, and only the records of scientists experienced with the species and the individuals concerned should be used. Even then, interpretation of a single record is always ambiguous, and analysis is feasible only when collation of multiple records shows that a behaviour pattern occurs repeatedly under similar circumstances. This approach has been used successfully to study a number of creative capacities of animals: the distribution, nature and neural correlates of deception across the primate order; the occurrence of teaching in animals; and the neural correlates of several aptitudes-in birds, foraging innovation, and in primates, innovation, social learning and tool-use. Drawing on these approaches, we describe the use of this method to investigate a new problem, the cognition of the African elephant, a species whose sheer size and evolutionary distance from humans renders the conventional methods of comparative psychology of little use. The aim is both to chart the creative cognitive capacities of this species, and to devise appropriate experimental methods to confirm and extend previous findings.

Bradshaw, G.A., Schore, A.N., 2007. How elephants are opening doors: Developmental neuroethology, attachment and social context. Ethology 113, 426-436.
Abstract: Ethology's renewed interest in developmental context coincides with recent insights from neurobiology and psychology on early attachment. Attachment and social learning are understood as fundamental mechanisms in development that shape core processes responsible for informing behaviour throughout a lifetime. Each field uniquely contributes to the creation of an integrated model and encourages dialogue between Tinbergen's four analytical levels: ethology in its underscoring of social systems of behaviour and context, psychology in its emphasis on socio-affective attachment transactions, and neuroscience in its explication of the coupled development of brain and behaviour. We review the relationship between developmental context and behaviour outcome as a topic shared by the three disciplines, with a specific focus on underlying neuroethological mechanisms. This interdisciplinary convergence is illustrated through the example of abnormal behaviour in wild African elephants (Loxodonta africana) that has been systematically observed in human-caused altered social contexts. Such disruptions impair normative socially mediated neuroendocrinological development leading to psychobiological dysregulation that expresses as non-normative behaviour. Aberrant behaviour in wild elephants provides a critical field example of what has been established in ex situ and clinical studies but has been largely absent in wild populations: a concrete link between effects of human disturbance on social context, and short- and long-term neuroethology. By so doing, it brings attention to the significant change in theories of behaviour that has been occurring across disciplines - namely, the merging of psychobiological and ethological perspectives into common, cross-species, human inclusive models.

King, L.E., Douglas-Hamilton, I., Vollrath, F., 2007. African elephants run from the sound of disturbed bees. Current Biology 17, 832-833.
Abstract: Encroaching human development into former wildlife areas [1] is compressing African elephants into ever smaller home ranges, causing increased levels of human-elephant conflict [2]. African honeybees have been proposed as a possible deterrent to elephants [3]. We have performed a sound playback experiment to study this hypothesis. We found that a significant majority of elephants, in a sample of 18 well-known families and subgroups of varying sizes, reacted negatively - immediately walking or running away - when they heard the buzz of disturbed bees, while they ignored the control sound of natural white-noise. Whether the observed response was the result of individual conditioning or of learning by social facilitation remains to be established. Our study strongly supports the hypothesis that bees - and perhaps even their buzz alone - may be deployed to keep elephants at bay.

Maple, T.L., 2007. Toward a science of welfare for animals in the zoo. J. Appl. Anim Welf. Sci. 10, 63-70.
Abstract: Although the accredited institutions of the Association of Zoos and Aquariums have all committed to enhancing the welfare of nonhuman animals, acceptable standards and best practices are still under debate. Currently, experts from zoos and the field hold widely divergent opinions about exhibition and management standards for elephants. Standards and practices for managing nonhuman primates provide a model for other nonhuman creatures exhibited in zoos and aquariums. Examining the key issues for primates demonstrates the value of applying scientific data before promulgating standards. The field of applied behavior analysis provides a wealth of information to frame the debate. Animal behaviorists have contributed to an emerging science of animal welfare, which may provide a foundation for empirical zoo management, standards, and practices

Maple, T.L., 2007. Toward a science of welfare for animals in the zoo. J Appl Anim Welf Sci 10, 63-70.
Abstract: Although the accredited institutions of the Association of Zoos and Aquariums have all committed to enhancing the welfare of nonhuman animals, acceptable standards and best practices are still under debate. Currently, experts from zoos and the field hold widely divergent opinions about exhibition and management standards for elephants. Standards and practices for managing nonhuman primates provide a model for other nonhuman creatures exhibited in zoos and aquariums. Examining the key issues for primates demonstrates the value of applying scientific data before promulgating standards. The field of applied behavior analysis provides a wealth of information to frame the debate. Animal behaviorists have contributed to an emerging science of animal welfare, which may provide a foundation for empirical zoo management, standards, and practices.

Meller, C.L., Croney, C.C., Shepherdson, D., 2007. Effects of rubberized flooring on Asian elephant behavior in captivity. Zoo. Biol. 26, 51-61.
Abstract: Six Asian elephants at the Oregon Zoo were observed to determine the effects of a poured rubber flooring substrate on captive Asian elephant behavior. Room utilization also was evaluated in seven rooms used for indoor housing, including Front and Back observation areas. Data were collected in three phases. Phase I (Baseline Phase) examined elephant behavior on old concrete floors. In Phase II (Choice Phase), elephant behavior was observed in the Back observation area where room sizes were comparable and when a choice of flooring substrates was available. Phase III (Final Phase) examined elephant behavior when all rooms in both observation areas, Front and Back, were converted to rubberized flooring. Room use in both observation areas remained stable throughout the study, suggesting that flooring substrate did not affect room use choice. However, there was a clear pattern of decreased discomfort behaviors on the new rubber flooring. Normal locomotion as well as stereotypic locomotion increased on the new rubber flooring. In addition, resting behavior changed to more closely reflect the resting behavior of wild elephants, which typically sleep standing up, and spend very little time in lateral recumbence. Overall, these findings suggest that the rubber flooring may have provided a more comfortable surface for locomotion as well as standing resting behavior. It is suggested that poured rubber flooring may be a beneficial addition to similar animal facilities. Zoo Biol 0:1-11, 2007. (c) 2007 Wiley-Liss, Inc

Savage, V.M., West, G.B., 2007. A quantitative, theoretical framework for understanding mammalian sleep. Proc. Natl. Acad. Sci. U. S. A 104, 1051-1056.
Abstract: Sleep is one of the most noticeable and widespread phenomena occurring in multicellular animals. Nevertheless, no consensus for a theory of its origins has emerged. In particular, no explicit, quantitative theory exists that elucidates or distinguishes between the myriad hypotheses proposed for sleep. Here, we develop a general, quantitative theory for mammalian sleep that relates many of its fundamental parameters to metabolic rate and body size. Several mechanisms suggested for the function of sleep can be placed in this framework, e.g., cellular repair of damage caused by metabolic processes as well as cortical reorganization to process sensory input. Our theory leads to predictions for sleep time, sleep cycle time, and rapid eye movement time as functions of body and brain mass, and it explains, for example, why mice sleep approximately 14 hours per day relative to the 3.5 hours per day that elephants sleep. Data for 96 species of mammals, spanning six orders of magnitude in body size, are consistent with these predictions and provide strong evidence that time scales for sleep are set by the brain's, not the whole-body, metabolic rate

Stoeger-Horwath, A.S., Stoeger, S., Schwammer, H.M., Kratochvil, H., 2007. Call repertoire of infant African elephants: first insights into the early vocal ontogeny. J. Acoust. Soc. Am. 121, 3922-3931.
Abstract: African savannah elephants (Loxodonta africana) have a complex acoustic communication system, but very little is known about their vocal ontogeny. A first approach in ontogenetic studies is to define the call repertoire of specific age groups. Twelve hundred calls of 11 infant elephants from neonatal to 18 months of age recorded at the Vienna Zoo in Austria and at the Daphne Sheldrick's orphanage at the Nairobi National Park, Kenya were analyzed. Six call types were structurally distinguished: the rumble, the bark, the grunt, the roar (subdivided into a noisy-, tonal-, and mixed-roar), the snort, and the trumpet. Generally, within-call-type variation was high in all individuals. In contrast to adult elephants, the infants showed no gender-dependent variation in the structure or in the number of call types produced. Male infants, however, were more vocally adamant in their suckle behavior than females. These results give a first insight to the early vocal ontogeny and should promote further ontogenetic studies on elephants. Due to their vocal learning ability in combination with the complex fission-fusion society, elephants could be an interesting model to study the role of imitation in the vocal ontogeny of a nonprimate terrestrial mammal

Teixeira, C.P., Schetini de Azevedo, C., Mendl, M., Cipreste, C.F., Young, R.J., 2007. Revisiting translocation and reintroduction programmes: the importance of considering stress. Animal Behaviour 73,  1-13.
Abstract: It is widely known that the adverse effects of stress must be considered in animal conservation programmes. However, a full consideration of how and where stress occurs in animal conservation programmes has not been undertaken, especially in translocation and reintroduction programmes. The literature concerning these types of programmes shows high levels of mortality, despite researchers' consideration of the effects of stress. However, an analysis of the literature shows that many conservation biologists have only a superficial knowledge about stress. For example, most do not understand the importance of subclinical stress or the fact that the effect of successive stressors can be additive or accumulative. While most conservation biologists know that stress is bad for animal health, few have considered its adverse effects on cognitive abilities, which an animal needs to survive in the wild (e.g. memory). In this paper we conclude with suggestions for improving the efficiency of animal conservation programmes in terms of the number of animals surviving after reintroduction or translocation. The most important conclusion from this review of the literature is that there needs to be a greater interchange of information between animal welfare and animal conservation scientists.

Wittemyer, G., Ganswindt, A., Hodges, K., 2007. The impact of ecological variability on the reproductive endocrinology of wild female African elephants. Hormones and Behavior 51, 346-354.
Abstract: Non-invasive endocrine methods enable investigation of the relationship between ecological variation and ovarian activity and how this impacts on demographic processes. The underlying physiological factors driving high variation in inter-calving intervals among multi-parous African elephants offer an interesting system for such an investigation. This study investigates the relationship between Normalized Differential Vegetation Index (NDVI), an ecosystem surrogate measure of primary productivity, and fecal progestin concentrations among wild female elephants. Matched fecal samples and behavioral data on reproductive activity were collected from 37 focal individuals during the two-year study. Linear mixed models were used to explore the relationship between fecal 5 alpha-pregnane-3-ol-20-one concentrations and the independent variables of NDVI, calf sex, female age, gestation day, and time since last parturition. Among both non-pregnant and pregnant females, fecal 5 alpha-pregnane-3-ol-20-one concentrations were significantly correlated with time-specific NDVI indicating a strong relationship between ecological conditions and endocrine activity regulating reproduction. In addition, the age of a female and time since her last parturition impacted hormone concentrations. These results indicate that the identification of an individual's reproductive status from a single hormone sample is possible, but difficult to achieve in practice since numerous independent factors, particularly season, impact fecal hormone concentrations. Regardless of season, however, fecal 5 alpha-pregnane-3-ol-20-one concentrations below 1 mu g/g were exclusively collected from non-pregnant females, which could be used as a threshold value to identify non-pregnant individuals. Collectively the information generated contributes to a better understanding of environmental regulation of reproductive endocrinology in wild elephant populations, information salient to the management and manipulation of population dynamics in this species.

Wittemyer, G., Getz, W.M., 2007. Hierarchical dominance structure and social organization in African elephants, Loxodonta africana. Animal Behaviour 73, 671-681.
Abstract: According to the socioecological framework, transitivity (or linearity) in dominance relationships is related to competition over critical resources. When a population is structured into groups, the intensity of between- versus within-group competition influences the form and function of its social organization. Few studies have compared the type and relative intensity of competition at these two levels. African elephants have well-structured social relations, providing an exemplary system for such a study. We report on dominance hierarchies among free-ranging elephants and evaluate the factors that drive their socioecological structure to lie in a region of the three-dimensional nepotism/despotism/tolerance space rarely observed among social species; namely, where non-nepotistic, transitive dominance hierarchies within groups emerge despite kin-based philopatry and infrequent agonistic interactions over widely distributed resources. We found significant transitivity in dominance hierarchies between groups. Dominance relations among the matriarchs of different social groups were primarily age based, rather than driven by physical or group size, and group matriarch rank influenced the dominance relationships among nonmatriarchal females in the population. Our results suggest that between-group dominance relationships induce tolerance among group members, which in combination with high group relatedness, reduces the benefits of nepotism. We postulate that cognitive abilities and high risk of injury in contests enhance winner and loser effects, facilitating the formation of transitive dominance relationships, despite widely distributed resources over which infrequent competition occurs. The interplay of cognitive abilities, winner and loser effects, resource distribution, and within- and between-group dominance relationships may produce behaviour in other strongly social mammals that differs from that predicted by a superficial application of current socioecological models.

Alfa Gambari Imorou, S., Sinsin, B. Impact of elephant's behaviour on the dynamic of vegetation in the W-Regional Park: The case of elephants in the north of Benin. Proceedings International Elephant Conservation & Research Symposium.  227-240. 2006. 2006.
Ref Type: Conference Proceeding

Archie, E.A., Moss, C.J., Alberts, S.C., 2006. The ties that bind: genetic relatedness predicts the fission and fusion of social groups in wild African elephants
490. Proc. Biol. Sci. 273, 513-522.
Abstract: Many social animals live in stable groups. In contrast, African savannah elephants (Loxodonta africana) live in unusually fluid, fission-fusion societies. That is, 'core' social groups are composed of predictable sets of individuals; however, over the course of hours or days, these groups may temporarily divide and reunite, or they may fuse with other social groups to form much larger social units. Here, we test the hypothesis that genetic relatedness predicts patterns of group fission and fusion among wild, female African elephants. Our study of a single Kenyan population spans 236 individuals in 45 core social groups, genotyped at 11 microsatellite and one mitochondrial DNA (mtDNA) locus. We found that genetic relatedness predicted group fission; adult females remained with their first order maternal relatives when core groups fissioned temporarily. Relatedness also predicted temporary fusion between social groups; core groups were more likely to fuse with each other when the oldest females in each group were genetic relatives. Groups that shared mtDNA haplotypes were also significantly more likely to fuse than groups that did not share mtDNA. Our results suggest that associations between core social groups persist for decades after the original maternal kin have died. We discuss these results in the context of kin selection and its possible role in the evolution of elephant sociality

Bagley, K.R., Goodwin, T.E., Rasmussen, L.E.L., Schulte, B.A., 2006. Male African elephants, Loxodonta africana, can distinguish oestrous status via urinary signals. Animal Behaviour 71, 1445.
Abstract: African elephants are a polygynous species that raise offspring in a matriarchal society. Unlike females, males disperse, spend time in mate groups and search for mates when mature. Urinary chemical signals aid males in detecting reproductively active females. A preovulatory pheromone has been identified in Asian elephants, Elephas maximus, but has not yet been experimentally identified in African elephants. In this study, the goal was to determine whether adult captive male African elephants can distinguish between urine from conspecific females in luteal and periovulatory oestrous stages as an indication that a preovulatory pheromone is released in the urine. Urine was collected from seven different female African elephants during their luteal and periovulatory periods of oestrus. Bioassays were conducted with nine adult male elephants housed at six different facilities. Males were presented with the two urine types and a control sample once a day over 3 days to reduce sample novelty, which can result in misleadingly high responses. All mates showed greater chemosensory responses to the periovulatory urine by trial 3 with the ability to distinguish the urines increasing over the 3 days. This is the first experimental behavioural evidence that African elephants release an oestrous pheromone in the urine. The ability of the captive male elephants to discern between the two urine types bolsters the hypothesis that there is a preovulatory pheromone in African elephants and encourages efforts to identify it.

Bertschinger, H., Delsink, A., Kirkpatrick, J.F., Human, A., Grobler, D., van Altena, J.J. Management of elephant populations in private South African game reserves with porcine zona pellucida vaccine.  2006 Proceedings American Association of Zoo Veterinarians.  283-285. 2006.
Ref Type: Conference Proceeding
Abstract: Control of African elephant populations has become an absolute necessity in a number of game reserves in southern Africa.  The two main methods used to control populations so far are culling and translocation. Culling, besides being regarded as inhumane and unacceptable in many quarters, is not suitable for smaller populations.  It requires that whole family units are culled simultaneously which could mean that in reserves with 10 to 50 elephants a considerable portion, if not the entire population, is killed.  As far as translocation is concerned, limited new space is available for elephants. The only alternative to the two above options is to control the rate of reproduction. The porcine zona pellucida (pZP) vaccine has been used to successfully contracept wild horses and other wildlife species.  Work on the contraception of African elephants was initiated in the Kruger National Park in 1995 when the potential for using the porcine zona pellucida (pZP) was investigated. Subsequently the first field trials on wild elephants were carried out in Kruger and the results clearly showed that elephants could be contracepted with the pZP vaccine, although the efficacy achieved was 80%. During these field trials safety and reversibility werecould be demonstrated. In 2000 an elephant contraceptive program was initiated at Makalali Private Game Reserve, RSA, which has become the flagship model for immunocontrol in African elephants. The preliminary findings have been reported in three publications.During the first year, all 18 cows that were individually identified and older than 12 yr of age were treated.  During the next 4 yr the number of cows contracepted increased to 23 as young animals were added to the program. The standard vaccination procedure during the first year consisted of a primary vaccination (600 μg or 400 μg pZP with 0.5 ml Freund's modified complete adjuvant) followed by boosters (200 μg pZP with 0.5 ml Freund's incomplete adjuvant) at 3 to 6-wk intervals. Annual boosters to maintain antibody titers and contraceptive effect followed.  To date, the success rate on cows that have passed reserve-specific intercalving period of 56 mo has been 100%. The population stabilized within 3 yr by which time when all cows that had been pregnant at the time of first vaccination in 2000 had calved. Once again safety during pregnancy (14 cows pregnant at 2-21 mo gestation when first treated gave birth to normal healthy calves) as well as side effects that were limited to occasional lumps at the site of vaccination could be shown. Following ground darting, behavioral patterns returned to pre-darting status within 2 days. During 2003 and 2004 most boosters were administered from a helicopter; whereas, previously they had been done from a vehicle or on foot.  In all cases, drop-out darts were used. Time taken for vaccination from helicopter take-off to landing was about 30 min (1.5 min per cow; 30 min for total time). This required prior knowledge of the locations of family units or that an individual in each unit is radio-collared. Herds settled down much more quickly (1-2 days) than if darted from the ground. Since then we have vaccinated another 107 elephant cows in eight game reserves.  The cow populations have ranged from 4 to 43. In one of the reserves, Mabula, RSA, two of the four cows vaccinated have passed the mean intercalving intervals of the reserve with neither of them producing a calf. Treatment at the remaining reserves was initiated in 2004 or 2005 and it is too early to evaluate results.  The most difficult reserve in terms of the vaccination process was Welgevonden, RSA, (35 000 ha) with 43 cows.  The reserve is mountainous and heavily wooded. None of the elephants were collared and individuals could not be easily identified on the day of primary vaccination.  The total flying time during which individuals were identified and vaccinated was 4.5 hr.  Administration of the first booster took about 2 hr to locate and vaccinate each cow. Between the first and second booster the first rains occurred, followed by the spring flush of the vegetation. By the time the second booster was attempted late in November, the trees all had foliage. Only half the cows were located and darted because the elephants were very difficult to spot under the tree canopies.  The valuable lessons we learned from this were: 1) that helicopter vaccinations should be performed when most trees are bare, and 2) when larger populations are vaccinated repeatedly during the first year, one cow in each family unit should be radio-collared. This makes rapid location of each unit possible and cuts down on the major cost factor that is flying time. Elephant behavior is being monitored in all eight reserves where contraception is being applied. Because most of them have been contracepted recently, only the data from Makalali is available. The elephants at Makalali have been monitored intensively almost on a daily basis. To date, no anomalies in terms of aggressive or indifferent behavior with regards to nursing time, nursing behavior and calf proximity have been noted. No change in the cows' social hierarchy has been noted. Since January 2003, a total of 15 heats were observed in 10 cows (nine in 2003 and six in 2004) with four mating episodes. For the same period, 38 musth occasions were seen in five bulls (26 in 2003 and 12 in 2004). These occasions include musth displayed in the same bull during consecutive days or within the same musth cycle. The greatest occurrence of musth was recorded in the largest, dominant bull. Bulls were not observed harassing or separating cows off from their herds or calves as a result of increased estrous frequency. Thus, the Makalali program demonstrates that pZP does not cause herd fragmentation, harassment by bulls, change in rank and other negative behaviors normally associated with hormonal contraceptives. In conclusion we feel that it is important to emphasize the following points: The pZP vaccine can be used successfully to contracept African elephants The vaccine is safe during pregnancy and has no negative effect on birth or calf raising It has no side effects other than occasional swelling at the site of vaccination It is reversible Other than an increased incidence of heat no behavioral side effects were seen.

Cerling, T.E., Wittemyer, G., Rasmussen, H.B., Vollrath, F., Cerling, C.E., Robinson, T.J., Douglas-Hamilton, I., 2006. Stable isotopes in elephant hair document migration patterns and diet changes
521. Proc. Natl. Acad. Sci. U. S. A 103, 371-373.
Abstract: We use chronologies of stable isotopes measured from elephant (Loxodonta africana) hair to determine migration patterns and seasonal diet changes in elephants in and near Samburu National Reserve in northern Kenya. Stable carbon isotopes record diet changes, principally enabling differentiation between browse and tropical grasses, which use the C3 and C4 photosynthetic pathways, respectively; stable nitrogen isotopes record regional patterns related to aridity, offering insight into localized ranging behavior. Isotopically identified range shifts were corroborated by global positioning system radio tracking data of the studied individuals. Comparison of the stable isotope record in the hair of one migrant individual with that of a resident population shows important differences in feeding and ranging behavior over time. Our analysis indicates that differences are the result of excursions into mesic environments coupled with intermittent crop raiding by the migrant individual. Variation in diet, quantified by using stable isotopes, can offer insight into diet-related wildlife behavior

Clemins, P.J., Johnson, M.T., 2006. Generalized perceptual linear prediction features for animal vocalization analysis
431. J. Acoust. Soc. Am. 120, 527-534.
Abstract: A new feature extraction model, generalized perceptual linear prediction (gPLP), is developed to calculate a set of perceptually relevant features for digital signal analysis of animal vocalizations. The gPLP model is a generalized adaptation of the perceptual linear prediction model, popular in human speech processing, which incorporates perceptual information such as frequency warping and equal loudness normalization into the feature extraction process. Since such perceptual information is available for a number of animal species, this new approach integrates that information into a generalized model to extract perceptually relevant features for a particular species. To illustrate, qualitative and quantitative comparisons are made between the species-specific model, generalized perceptual linear prediction (gPLP), and the original PLP model using a set of vocalizations collected from captive African elephants (Loxodonta africana) and wild beluga whales (Delphinapterus leucas). The models that incorporate perceptional information outperform the original human-based models in both visualization and classification tasks

De, M.E., Cowlishaw, G., 2006. Species protection, the changing informal economy, and the politics of access to the bushmeat trade in the Democratic Republic of Congo
420. Conserv. Biol. 20, 1262-1271.
Abstract: Our understanding of the linkages between the bushmeat trade and the wider informal economy is limited. This lack of knowledge is particularly problematic for conservation under conditions of political instability, when the informal economy can be highly dynamic and impacts on wildlife populations can be severe. To explore these interlinked processes, we conducted a study of the bushmeat trade in Garamba National Park, Democratic Republic of Congo, through a combination of market surveys, semistructured interviews, and direct observation. We focused on the sale of protected and unprotected species in urban and rural markets, and the bushmeat commodity chains that supplied these markets, under conditions of political stability and armed conflict. During peacetime, protected species from the park (predominantly elephant and buffalo) rarely appeared in the rural markets, but they comprised more than half of all bushmeat sales in the urban markets. This pattern reflected differences in the rural and urban commodity chains. Automatic weapons were urban trade. The use of such weapons was discouraged by the traditional chiefs, who administered the village markets. During wartime, the sales of protected species in the urban markets increased fivefold because the military officers fled, leaving behind an open-access system that led to a massive increase in the exploitation of protected species. In contrast, the rural markets remained relatively stable because of the continued authority of the village chiefs. Our results indicate that sociopolitical factors can be an important determinant of species offtake and, therefore, that knowledge of the bushmeat commodity chain can be vital to controlling theprocesses that drive species extraction. In addition, our findings suggest that traditional authorities can be potentially valuable partners for bushmeat management

Druce, H., Pretorius, K., Druce, D., Slotow, R., 2006. The effect of mature elephant bull introductions on resident bull's group size and musth periods: Phinda Private Game Reserve, South Africa. South African Journal of Wildlife Research 36, 133-137.
Abstract: African elephants have been reintroduced into small, enclosed reserves in South Africa,many populations being established with orphans <10 years old.This has resulted in abnormal behaviour in some elephant populations, which was corrected in Pilanesberg National Park by introducing older bulls and culling certain problem elephants.In July 2003, three older bulls (29-41 years old) were introduced into Phinda Private Game Reserve, KwaZulu-Natal, South Africa, in order to normalize the bull age structure and in an attempt to reduce the abnormally long musth period of one particular resident bull. These introduced bulls were monitored intensively after release, as was the resident bull population, both before and after introduction of the older bulls.The introduced bulls all came into musth within eleven months postrelease.The older bulls do not appear to have had any influence on the musth periods of the oldest resident bull (36 years old at introduction). Detailed behavioural studies of the effects of management actions on elephant populations, within small, enclosed reserves provide information and resources for future management decisions.This study demonstrates that old bulls can be successfully introduced to very small areas provided that electrification of the entire perimeter is secure. Further, the introduction has no detectable medium-term (one year) effect on the behaviour of a relatively dense population of resident elephants, and the welfare of the elephants was not greatly affected.

Elzanowski, A., Sergiel, A., 2006. Stereotypic behavior of a female Asiatic elephant (Elephas maximus) in a zoo
381. J. Appl. Anim Welf. Sci. 9, 223-232.
Abstract: This study recorded daytime behavior of a female Asiatic elephant at the Municipal Zoo, Wroclaw, Poland, in both an indoor pen and an outdoor paddock as continuous scan sampling for 140 hr, over 35 days in 1 year. Stereotypic sequences involved bouts of highly repetitive stereotypic movements and much more variable interbout behavior. The study found both stereotypic movements, nodding and body (corpus) swaying, were asymmetric, accompanied by protraction of the right hind leg and to-and-fro swinging of the trunk. The elephant spent 52% of the daytime in stereotypic movements, 3.5 times the level reported for females in other zoos' groups. The share of time devoted to stereotypic behavior was lowest in the summer when the elephant was regularly released to the paddock and highest in the late fall after she had stayed in the pen after months of days outside. This suggests that changes in the management routine enhance stereotypies. Comparing the summer and winter stable management periods, stereotypies were much more frequent in the indoor pen than the outdoor paddock, suggesting that the confinement to a barren pen contributed to the observed levels of stereotypies

Evans, K. Adolescent male African elephant behaviour. Proceedings International Elephant Conservation & Research Symposium.  86. 2006. 2006.
Ref Type: Conference Proceeding

Kandler, C., Schwammer, H., Becker, M., Fleissner, G. Chronoethology of African elephants (Loxodonta africana) in Zoological Gardens. Proceedings International Elephant Conservation & Research Symposium.  93-97. 2006. 2006.
Ref Type: Conference Proceeding

Langbauer, W., Philp, K., Frydman, G., Galvanek, J. The effect of human contact on African elephant heart rate. Proceedings International Elephant Conservation & Research Symposium.  253-255. 2006. 2006.
Ref Type: Conference Proceeding

Larke, A., Crews, D.E., 2006. Parental investment, late reproduction, and increased reserve capacity are associated with longevity in humans. J Physiol Anthropol 25, 119-131.
Abstract: Throughout the living world trade-offs between reproductive success and longevity have been observed. In general, two extremes of life history patterning are reported, r- and K-selected species. The latter tend toward larger body sizes, few offspring from any one pregnancy, few offspring over the female reproductive span, longer life spans, and greater parental investment (PI: all efforts and expenses associated with the production, gestation, post-natal care, feeding, and protection of young) (e.g., whales, elephants, hominids). r-selected species tend toward smaller body size, multiple births/litters per pregnancy, female production of many gametes and offspring over the life span, and low levels of PI (e.g., most plants, insects, mice). These differences have significant influences on physiological variation among human populations.Across human samples, reproductive success (RS: the number of offspring successfully birthed and reared to reproductive age) has been reported to vary positively, negatively, and not at all with longevity of women. This complexity may be in part due to the fact that both early-life and late-life fecundity are associated with longevity in women, while total parity seems a poor gauge of female longevity in humankind. Large variations in associations of RS with longevity in women suggest that multiple factors may confound this association. One confounding factor is that among women, RS is largely determined not by fecundity, but by the quality of PI available to offspring. Among modern humans, PI is more complex, longer lasting (both relatively and absolutely), and extensive than for any other ammal. This suggests that modern human life history is a reflection of the co-evolution of longevity and extensive PI as part of our species' biocultural evolution. The need for long-term PI has greatly shaped human physiological variation and patterns of longevity.

Liang, Y., McMeeking, R.M., Evans, A.G., 2006. A finite element simulation scheme for biological muscular hydrostats
478. Journal of Theoretical Biology 242, 142-150.
Abstract: An explicit finite element scheme is developed for biological muscular hydrostats such as squid tentacles, octopus arms and elephant trunks. The scheme is implemented by embedding muscle fibers in finite elements. In any given element, the fiber orientation can be assigned arbitrarily and multiple muscle directions can be simulated. The mechanical stress in each muscle fiber is the sum of active and passive parts. The active stress is taken to be a function of activation state, muscle fiber shortening velocity and fiber strain; while the passive stress depends only on the strain. This scheme is tested by simulating extension of a squid tentacle during prey capture; our numerical predictions are in close correspondence with existing experimental results. It is shown that the present finite element scheme can successfully simulate more complex behaviors such as torsion of a squid tentacle and the bending behavior of octopus arms or elephant trunks

Morris, S., Humphreys, D., Reynolds, D., 2006. Myth, marula, and elephant: an assessment of voluntary ethanol intoxication of the African elephant (Loxodonta africana) following feeding on the fruit of the marula tree (Sclerocarya birrea)
485. Physiol Biochem. Zool. 79, 363-369.
Abstract: Africa can stir wild and fanciful notions in the casual visitor; one of these is the tale of inebriated wild elephants. The suggestion that the African elephant (Loxodonta africana) becomes intoxicated from eating the fruit of the marula tree (Sclerocarya birrea) is an attractive, established, and persistent tale. This idea now permeates the African tourist industry, historical travelogues, the popular press, and even scholastic works. Accounts of ethanol inebriation in animals under natural conditions appear mired in folklore. Elephants are attracted to alcohol, but there is no clear evidence of inebriation in the field. Extrapolating from human physiology, a 3,000-kg elephant would require the ingestion of between 10 and 27 L of 7% ethanol in a short period to overtly affect behavior, which is unlikely in the wild. Interpolating from ecological circumstances and assuming rather unrealistically that marula fruit contain 3% ethanol, an elephant feeding normally might attain an ethanol dose of 0.3 g kg(-1), about half that required. Physiological issues to resolve include alcohol dehydrogenase activity and ethanol clearance rates in elephants, as well as values for marula fruit alcohol content. These models were highly biased in favor of inebriation but even so failed to show that elephants can ordinarily become drunk. Such tales, it seems, may result from "humanizing" elephant behavior

Nissani, M., 2006. Do Asian elephants (Elephas maximus) apply causal reasoning to tool-use tasks?
511. J. Exp. Psychol. Anim Behav. Process 32, 91-96.
Abstract: Two experiments addressed contradictory claims about causal reasoning in elephants. In Experiment 1, 4 Asian elephants (Elephas maximus) were pretrained to remove a lid from the top of a bucket and retrieve a food reward. Subsequently, in the first 5 critical trials, when the lid was placed alongside the bucket and no longer obstructed access to the reward, each elephant continued to remove the lid before retrieving the reward. Experiment 2, which involved 11 additional elephants and variations of the original design, yielded similarly counterintuitive observations. Although the results are open to alternative interpretations, they appear more consistent with associative learning than with causal reasoning. Future applications of Fabrean methodologies (J. H. Fabre, 1915) to animal cognition are proposed

Norgaard, C. Towards 24 hours of enrichment. Proceedings International Elephant Conservation & Research Symposium.  260. 2006. 2006.
Ref Type: Conference Proceeding

Plotnik, J.M., de Waal, F.B., Reiss, D., 2006. Self-recognition in an Asian elephant
389. Proc. Natl. Acad. Sci. U. S. A 103, 17053-17057.
Abstract: Considered an indicator of self-awareness, mirror self-recognition (MSR) has long seemed limited to humans and apes. In both phylogeny and human ontogeny, MSR is thought to correlate with higher forms of empathy and altruistic behavior. Apart from humans and apes, dolphins and elephants are also known for such capacities. After the recent discovery of MSR in dolphins (Tursiops truncatus), elephants thus were the next logical candidate species. We exposed three Asian elephants (Elephas maximus) to a large mirror to investigate their responses. Animals that possess MSR typically progress through four stages of behavior when facing a mirror: (i) social responses, (ii) physical inspection (e.g., looking behind the mirror), (iii) repetitive mirror-testing behavior, and (iv) realization of seeing themselves. Visible marks and invisible sham-marks were applied to the elephants' heads to test whether they would pass the litmus "mark test" for MSR in which an individual spontaneously uses a mirror to touch an otherwise imperceptible mark on its own body. Here, we report a successful MSR elephant study and report striking parallels in the progression of responses to mirrors among apes, dolphins, and elephants. These parallels suggest convergent cognitive evolution most likely related to complex sociality and cooperation

Reznikova, Z., 2006. [The study of tool use as the way for general estimation of cognitive abilities in animals]
496. Zh. Obshch. Biol. 67, 3-22.
Abstract: Investigation of tool use is an effective way to determine cognitive abilities of animals. This approach raises hypotheses, which delineate limits of animal's competence in understanding of objects properties and interrelations and the influence of individual and social experience on their behaviour. On the basis of brief review of different models of manipulation with objects and tools manufacturing (detaching, subtracting and reshaping) by various animals (from elephants to ants) in natural conditions the experimental data concerning tool usage was considered. Tool behaviour of anumals could be observed rarely and its distribution among different taxons is rather odd. Recent studies have revealed that some species (for instance, bonobos and tamarins) which didn't manipulate tools in wild life appears to be an advanced tool users and even manufacturers in laboratory. Experimental studies of animals tool use include investigation of their ability to use objects physical properties, to categorize objects involved in tool activity by its functional properties, to take forces affecting objects into account, as well as their capacity of planning their actions. The crucial question is whether animals can abstract general principles of relations between objects regardless of the exact circumstances, or they develop specific associations between concerete things and situations. Effectiveness of laboratory methods is estimated in the review basing on comparative studies of tool behaviour, such as "support problem", "stick problem", "tube- and tube-trap problem", and "reserve tube problem". Levels of social learning, the role of imprinting, and species-specific predisposition to formation of specific domains are discussed. Experimental investigation of tool use allows estimation of the individuals' intelligence in populations. A hypothesis suggesting that strong predisposition to formation of specific associations can serve as a driving force and at the same time as obstacle to animals' activity is discussed. In several "technically gifted" species (such as woodpecker finches, New Caledonian crows, and chimpanzees) tool use seems to be guided by a rapid process of trial and error learning. Individuals that are predisposed to learn specific connections do this too quickly and thus become enslaved by stereotypic solutions of raising problems.

Shannon, G., Page, B.R., Duffy, K.J., Slotow, R., 2006. The role of foraging behaviour in the sexual segregation of the African elephant. Oecologia Epub.
Abstract: Elephants (Loxodonta africana) exhibit pronounced sexual dimorphism, and in this study we test the prediction that the differences in body size and sociality are significant enough to drive divergent foraging strategies and ultimately sexual segregation. Body size influences the foraging behaviour of herbivores through the differential scaling coefficients of metabolism and gut size, with larger bodied individuals being able to tolerate greater quantities of low-quality, fibrous vegetation, whilst having lower mass-specific energy requirements. We test two distinct theories: the scramble competition hypothesis (SCH) and the forage selection hypothesis (FSH). Comprehensive behavioural data were collected from the Pongola Game Reserve and the Phinda Private Game Reserve in South Africa over a 2.5-year period. The data were analysed using sex as the independent variable. Adult females targeted a wider range of species, adopted a more selective foraging approach and exhibited greater bite rates as predicted by the body size hypothesis and the increased demands of reproductive investment (lactation and pregnancy). Males had longer feeding bouts, displayed significantly more destructive behaviour (31% of observations, 11% for females) and ingested greater quantities of forage during each feeding bout. The independent ranging behaviour of adult males enables them to have longer foraging bouts as they experience fewer social constraints than females. The SCH was rejected as a cause of sexual segregation due to the relative abundance of low quality forage, and the fact that feeding heights were similar for both males and females. However, we conclude that the differences in the foraging strategies of the sexes are sufficient to cause spatial segregation as postulated by the FSH. Sexual dimorphism and the associated behavioural differences have important implications for the management and conservation of elephant and other dimorphic species, with the sexes effectively acting as distinct "ecological species".

Shannon, G., Page, B.R., Duffy, K.J., Slotow, R., 2006. The consequences of body size dimorphism: Are African elephants sexually segregated at the habitat scale? Behaviour 143, 1145-1168.
Abstract: Sexual segregation is a commonly observed phenomenon in dimorphic ungulates, which has been categorised into two distinct components: social segregation and habitat segregation. In this study we investigated whether elephants were sexually segregated at the habitat scale. The locations of 12 family groups and 16 males, in three distinct populations were recorded over a period of 2.5 years. Selection ratios were calculated for each habitat type and a Kendall's coefficient of concordance was used for the analyses. The habitat and foraging preferences were firstly tested for concordance within sex, and then between the sexes. Female habitat preferences showed significant concordance across all reserves and they also exhibited strong concordance in their summer foraging preferences. Their weakest association with habitat and foraging preference was during winter, which may be related to resource scarcity. Males exhibited significant concordance in their habitat preferences in two out of the three reserves. They had their weakest associations in the summer months and this may be linked to avoidance of other males in musth and the abundance of forage. There were no significant differences in habitat preference between males and females and it is likely that individual preferences vary as much within sex as between sexes. Differential habitat utilisation does not appear to be driving sexual segregation in elephants and it is postulated that sociality, divergent reproductive strategies and foraging behaviour at the plant scale play a more significant role. The results of this study highlight the importance of scale in elucidating the mechanisms involved in sexual segregation.

Shoshani, J., Kupsky, W.J., Marchant, G.H., 2006. Elephant brain. Part I: gross morphology, functions, comparative anatomy, and evolution. Brain Res Bull 70, 124-157.
Abstract: We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior.

Shoshani, J., Kupsky, W.J., Marchant, G.H., 2006. Elephant brain. Part I: gross morphology, functions, comparative anatomy, and evolution
446. Brain Res. Bull. 70, 124-157.
Abstract: We report morphological data on brains of four African, Loxodonta africana, and three Asian elephants, Elephas maximus, and compare findings to literature. Brains exhibit a gyral pattern more complex and with more numerous gyri than in primates, humans included, and in carnivores, but less complex than in cetaceans. Cerebral frontal, parietal, temporal, limbic, and insular lobes are well developed, whereas the occipital lobe is relatively small. The insula is not as opercularized as in man. The temporal lobe is disproportionately large and expands laterally. Humans and elephants have three parallel temporal gyri: superior, middle, and inferior. Hippocampal sizes in elephants and humans are comparable, but proportionally smaller in elephant. A possible carotid rete was observed at the base of the brain. Brain size appears to be related to body size, ecology, sociality, and longevity. Elephant adult brain averages 4783 g, the largest among living and extinct terrestrial mammals; elephant neonate brain averages 50% of its adult brain weight (25% in humans). Cerebellar weight averages 18.6% of brain (1.8 times larger than in humans). During evolution, encephalization quotient has increased by 10-fold (0.2 for extinct Moeritherium, approximately 2.0 for extant elephants). We present 20 figures of the elephant brain, 16 of which contain new material. Similarities between human and elephant brains could be due to convergent evolution; both display mosaic characters and are highly derived mammals. Humans and elephants use and make tools and show a range of complex learning skills and behaviors. In elephants, the large amount of cerebral cortex, especially in the temporal lobe, and the well-developed olfactory system, structures associated with complex learning and behavioral functions in humans, may provide the substrate for such complex skills and behavior

Stoeger-Horwath, A.S., Schwammer, H.M., Kratochvil, H., Stoeger, S. Infant talk - first insights into the vocal ontogeny of elephants (Loxodonta africana). Proceedings International Elephant Conservation & Research Symposium.  178-181. 2006. 2006.
Ref Type: Conference Proceeding

Tresz, H., 2006. Behavioral management at the Phoenix Zoo: New strategies and perspectives. Journal of Applied Animal Welfare Science 9, 65-70.
Abstract: It all started with a seemingly simple decision to re-evaluate and document the Phoenix Zoo's behavioral management protocol. The purpose of this project was to present proactive standards for the care and psychological well-being of our living collection, while meeting or exceeding the guidelines of the Animal Welfare Act. Preparing the protocol was a catalyst to re-evaluate the zoo's philosophy and application of behavioral management. It suggested a restructuring of collection management and the rethinking of future goals and practices. Gradually, the process became more focused and organized. Behavioral enrichment, training, animal behavior issues, and exhibit architecture were embraced as essential components for providing quality of life. Staff from all levels worked side-by-side on assignments. Our way of thinking and working was changing.

Vidya, T.N.C., Sukumar, R. Social organization of the Asian elephant (Elephas maximus) in southern India as inferred from microsatellite DNA. Proceedings International Elephant Conservation & Research Symposium.  214-216. 2006. 2006.
Ref Type: Conference Proceeding

Viljoen, J.J., Reynecke, H.C., du Toit, J.T., Langbauer, W.R. Elephant family groups may cause little environmental damage in the Kruger Park. Proceedings International Elephant Conservation & Research Symposium.  274. 2006. 2006.
Ref Type: Conference Proceeding

Weidner, E.B., Isaza, R., Galle, L.E., Barrie, K., Lindsay, W.A., 2006. Medical management of a corneal stromal abscess in a female Asisan elephant (Elephas maximus). Journal of Zoo and Wildlife Medicine 37, 397-400.

Weissenböck, N.M. How do elephants deal with various climate conditions? Previous results, recent data and new hypotheses. Proceedings International Elephant Conservation & Research Symposium.  217-224. 2006. 2006.
Ref Type: Conference Proceeding

Wittemyer, G., Getz, W.M., 2006. A likely ranking interpolation for resolving dominance orders in systems with unknown relationships. Behaviour 143, 909-930.
Abstract: In many animal systems agonistic interactions may be rare or not overt, particularly where such interactions are costly or of high risk as is common for large mammals. We present a technique developed specifically for resolving an optimized dominance order of individuals in systems with transitive (i.e. linear) dominance relationships, but where not all relationships are known. Our method augments the widely used I&SI method (de Vries, 1998) with an interpolation function for resolving the relative ranks of individuals with unknown relationships. Our method offers several advantages over other dominance methods by enabling the incorporation of any proportion of unknown relationships, resolving a unique solution to any dominance matrix, and calculating cardinal dominance strengths for each individual. As such, this method enables novel insight into difficult to study behavioural systems.

Bertelsen, M.F., Bojesen, M., Olsen, K.E.P. Fatal enterocolitis in two Asian elephants (Elephas maximus) caused by Clostridium difficile. 2005 Proceedings AAZV, AAWV, AZA Nutrition Advisory Group.  66-67. 2005.
Ref Type: Conference Proceeding
Abstract: Altered behavior, anorexia and listlessness were observed in four of five adult captive female Asian elephants (Elephas maximus). Two animals recovered, while two died after 2 days. The dead elephants were subjected to post mortem examination including histopathology, demonstrating fibrinonecrotic enteritis and colitis. Clostridium difficile was isolated from both dead elephants and from the feces of the two surviving affected animals, and identified by selective cultivation and PCR identification. All isolates had the tcdA and tcdB toxin genes and were positive in a toxigenic culture assay. C. difficile toxin from the intestinal content of one of the fatal cases was demonstrated using cell-culture based cytotoxin assays. Clostridium perfringens type A and Clostridium septicum were also isolated from both dead animals. Although C. perfringens has been associated with ulcerative enteritis in an elephant,¹ in this case these isolates likely are incidental, as C. perfringens enterotoxin was not demonstrated, and as C. septicum is well known for producing rapid post mortem overgrowth.  Amplified fragment length polymorphism typing, showed that the C. difficile isolates recovered from the outbreak, all had the same fingerprint profile, indicating that all four elephants were affected by the same bacterial clone. These findings appear to be the first to demonstrate that C. difficile may cause enterocolitis in elephants. The results emphasize the need to regard this organism as potentially dangerous for elephants. Although there was no prior exposure to antibiotic agents in this case, caution is recommended when treating elephants with antibiotics, as this may trigger C. difficile induced enterocolitis in other species, most notably humans and horses.²
LITERATURE CITED
1 Bacciarini, L.N., O. Pagan, J. Frey, and A. Grone. 2001. Clostridium perfringens beta2-toxin in an African elephant (Loxodonta africana) with ulcerative enteritis. Vet. Rec. 149: 618-20.
2 Songer, J.G. 1996. Clostridial enteric diseases of domestic animals. Clin. Microbiol. Rev. 9: 216-234.

Bradshaw, G.A., Schore, A.N., Brown, J.L., Poole, J.H., Moss, C.J., 2005. Elephant breakdown
639. Nature 433, 807.

Clemins, P.J., Johnson, M.T., Leong, K.M., Savage, A., 2005. Automatic classification and speaker identification of African elephant (Loxodonta africana) vocalizations
633. J. Acoust. Soc. Am. 117, 956-963.
Abstract: A hidden Markov model (HMM) system is presented for automatically classifying African elephant vocalizations. The development of the system is motivated by successful models from human speech analysis and recognition. Classification features include frequency-shifted Mel-frequency cepstral coefficients (MFCCs) and log energy, spectrally motivated features which are commonly used in human speech processing. Experiments, including vocalization type classification and speaker identification, are performed on vocalizations collected from captive elephants in a naturalistic environment. The system classified vocalizations with accuracies of 94.3% and 82.5% for type classification and speaker identification classification experiments, respectively. Classification accuracy, statistical significance tests on the model parameters, and qualitative analysis support the effectiveness and robustness of this approach for vocalization analysis in nonhuman species

Clemins, P.J., Johnson, M.T., Leong, K.M., Savage, A., 2005. Automatic classification and speaker identification of African elephant (Loxodonta africana) vocalizations. J Acoust Soc Am. 117, 956-963.
Abstract: A hidden Markov model (HMM) system is presented for automatically classifying African elephant vocalizations. The development of the system is motivated by
successful models from human speech analysis and recognition. Classification features include frequency-shifted Mel-frequency cepstral coefficients (MFCCs)
and log energy, spectrally motivated features which are commonly used in human speech processing. Experiments, including vocalization type classification and
speaker identification, are performed on vocalizations collected from captive elephants in a naturalistic environment. The system classified vocalizations with accuracies of 94.3% and 82.5% for type classification and speaker identification classification experiments, respectively. Classification accuracy, statistical significance tests on the model parameters, and qualitative analysis support the effectiveness and robustness of this approach for vocalization analysis in nonhuman species.
Speech and Signal Processing Laboratory, Marquette University, Milwaukee, Wisconsin 53233-1881, USA. patrick.clemins@marquette.edu

Cushman, S.A., Chase, M., Griffin, C., 2005. Elephants in space and time. Oikoso 109, 331-341.
Abstract: Autocorrelation in animal movements can be both a serious nuisance to analysis and a source of valuable information about the scale and patterns of animal behavior, depending on the question and the techniques employed. In this paper we present an approach to analyzing the patterns of autocorrelation in animal movements that provides a detailed picture of seasonal variability in the scale and patterns of movement. We used a combination of moving window Mantel correlograms, surface correlation and crosscorrelation analysis to investigate the scales and patterns of autocorrelation in the movements of three herds of elephants in northern Botswana. Patterns of autocorrelation of elephant movements were long-range, temporally complicated, seasonally variable, and closely linked with the onset of rainfall events. Specifically, for the three elephant herds monitored there was often significant autocorrelation among locations up to lags of 30 days or more. During many seasonal periods there was no indication of decreasing autocorrelation with increasing time between locations. Over the course of the year, herds showed highly variable and complex patterns of autocorrelation, ranging from random use of temporary home ranges, periodic use of focal areas, and directional migration. Even though the patterns of autocorrelation were variable in time and quite complex, there were highly significant correlations among the autocorrelation patterns of the different herds, indicating that they exhibited similar patterns of movement through the year. These major patterns of autocorrelation seem to be related to patterns of rainfall. The strength of correlation in movement patterns of the different herds decreased markedly at the cessation of major rain events. Also, there was a strong crosscorrelation between strength of autocorrelation of movement and rainfall, peaking at time lags of between three and four weeks. Overall, these approaches provide a powerful way to explore the scales and patterns of autocorrelation of animal movements, and to explicitly link those patterns to temporally variable environmental attributes, such as rainfall or vegetation phenology.

Deem, S.L., Brown, J.L., Eggert, L., Wemmer, C., Htun, W., Nyunt, T., Murray, S., Leimgruber, P. Health and management of working elephants in Myanmar (Burma). Procedings American Association of Zoo Veterinarians.  228-231. 2005.
Ref Type: Conference Proceeding
Abstract: Myanmar has approximately 6,000 working elephants.  Remaining wild elephants are declining, partly because of live-capture for captivity.  Through health and reproductive assessments, genetic analyses and GPS tracking of captive and wild elephants, we are exploring linkages between the two populations and conducting studies to reduce morbidity and mortality of captive elephants. Captive elephants live and work in Myanmar's forests in close proximity and contact to the remaining wild herds. We propose that reducing morbidity and mortality in the captive elephants will decrease the need for live-capture, and the risk of disease transmission, to wild elephants.
Introduction
There are an estimated 6,000 working elephants in Myanmar - half owned by the government operated Myanmar Timber Enterprise (MTE) and half owned privately^.5 This may be one of the largest captive elephant populations in the world and its management will have a significant impact on remaining wild herds in Myanmar.^4,6,8  With mortality rates higher than birth rates, the working population is probably maintained by supplementing it with elephants captured from the wild.⁵ There is evidence that continued harvest of wild elephants may have reduced the remaining wild populations of Myanmar.  Recent surveys of wild populations in two of Myanmar's protected elephant ranges revealed extremely low dung counts, indicative of small and declining herds. Constant contact with captive elephants in Myanmar's forests may exacerbate the threat to Myanmar wild elephants by increasing the transmission of disease between these two groups. For both the above reasons, we believe that the conservation of wild elephants in Myanmar will require significant improvements in the care and management of currently existing captive populations.  
Elephants owned by MTE receive veterinary care from the Burmese veterinarians that work for the timber company and travel extensively throughout the country to sites were the elephants are located.¹ There is a dire need for veterinary supplies and laboratory capabilities in the country. Currently, veterinary practices are based on the extensive field experience of lead MTE veterinarians. However, MTE veterinarians frequently rely on older published work ^3,7 and would benefit significantly from training that incorporates new insights into elephant health and new veterinary techniques. Similarly, because of their close-up experience of elephant health problems in the forests, MTE veterinarians may be able to make important new contributions to the care and management of elephants elsewhere.     
The overall objective of our study is to work jointly with MTE veterinarians to develop long-term captive population management strategies to reduce mortality and increase births in the working timber elephants and stop the continued off-take of animals from the wild to supplement captive herds.
Methods
The health component of this study has five major objectives.  These are to:
1              Conduct a training workshop, in conjunction with MTE veterinarians, on elephant management and veterinary care. 
2              Develop protocols so that the MTE veterinarians can collect samples for reproductive, genetic, and health status assessments.
3              Analyze samples and provide data to MTE veterinarians to improve husbandry, preventive care and disease treatment of working elephants.
4              Develop a comprehensive bibliography of all published information on the health and management of Myanmar elephants.
5              Perform an epidemiologic evaluation of records available on the historic and current working elephant population.
Specific steps to achieve these objectives include: 
1              Determine causes and rates of morbidity and mortality of captive MTE elephants.
2              Determine causes of low rates of reproduction in captivity.
3              Develop a genetic profile of the captive herds.
4              Develop a protocol to assess oozies-Burmese mahout-expertise in parallel with endocrine and health assessments to determine quality of care and potentially related stress.
5              Develop small population viability models to assess how current mortality effects long-term survival of the captive population and what supplementation from the wild is needed for short- and long-term sustainability.
6              Use population viability models to demonstrate how supplementation from the wild will negatively affect that population.
7              Get baseline health parameter data on free-ranging elephants.
8              Quantify habitat/space use using GPS and satellite tracking of captive and wild elephants. 
Results and Discussion
During an initial exploratory visit in November 2004, we learned that the annual mortality rate for MTE working elephants was 2.4% (66) in 2003.  Deaths occurred in all age groups (>18 yr, n = 40; 4 - 17 yr, n = 11; <4 yr, n = 15) and included preventable diseases (i.e., poor nutrition, heat stroke, diarrhea, dystocia, infectious and parasitic agents).  Additionally, we collected samples for performing health, genetic and endocrine analyses of 22 elephants maintained in one of the working camps (results to be presented). A relationship also was established with the veterinary staff at the Yangon Zoo, including follow up donations of veterinary literature and journals to the zoo. We provided medical advice for the care of an orphaned elephant calf and other animals housed at the zoo during our brief visit. We are seeking funds for a training course to be conducted in late 2005 and hope to perform health evaluations on a larger number of zoo and working elephants during that visit.
The National Zoo already has an extensive conservation program for wild elephants in Myanmar.^4,6,8  This program has focused on assessing wild elephant populations in protected areas and satellite-tracking of four wild elephants to learn more about their conservation status and ecology in Myanmar.  Currently this work is being extended to a national elephant survey. Part of this work included collecting fecal samples for genetic and health assessments.
The Smithsonian team of researchers involved in this project includes a veterinarian, reproduction physiologist, geneticist, conservation biologist, and landscape ecologist.  All members of this multidisciplinary team have extensive experience working with elephants and together provide the necessary expertise to study and understand the numerous factors affecting Myanmar's captive elephants and the long-term survival of elephants in Myanmar.  These challenges range from human land use and elephant population fragmentation, human-elephant conflict, poor reproduction and health care of captive elephants and lack of information on the health status of the wild elephants.  A viable conservation initiative for the elephants of Myanmar requires that health issues be addressed as one component of a comprehensive program to address the anthropogenic pressures on both working and wild elephants.²
The elephants of Myanmar are an excellent example of the fine line that exists between captive and wild animals, especially as it relates to health.  Captive and wild elephants are regularly in direct and indirect contact.  The working elephants live with their oozies who may expose them to diseases, such as tuberculosis.  The working elephants in turn may encounter wild elephants at night in the forests where they forage and live during non-working hours. In fact, the majority of captive born calves are said to be sired by wild bulls.  Potentially, the use of working elephants in selectively extracting valuable timber provides new strategies for the conservation of elephants and forests. Most likely, "elephant-logging" is less damaging than machine-operated timbering projects that tend to clear-cut areas and also damage the soil and streams.  However, decreasing the negative impact of such practices (i.e., minimizing off-take of elephants from the wild, decreasing disease risks to the wild elephants) is imperative.  
LITERATURE CITED
1 Aung, T., and T. Nyunt.  2002.  The care and management of the domesticated Asian elephant in Myanmar.  In: Baker, I., and M. Kashio (eds.): Giants on our hands. Proc. Int. Workshop Domesticated Asian Elephant. Dharmasarn Co., Ltd. Bangkok, Thailand. Pp. 89 - 102.
2 Deem, S.L., W.B. Karesh, and W. Weisman.  2001.  Putting theory into practice: wildlife health in conservation.  Conserv. Biol. 5: 1224-1233.
3 Evans, G.H. 1910.  Elephants and Their Diseases.  Government Printing. Rangoon. 323 
4 Kelly, D.S. 2005.  Habitat selection in declining elephant populations of Alaungdaw Kathapa National Park. Masters Thesis.  George Mason University.
5 Lair, R.C. 1997.  Myanmar. In: Gone Astray: The Care and Management of the Asian Elephant in Domesticity. FAO Regional Office for Asia and the Pacific, Thailand.  RAP Publication. Pp. 99-131
6 Leimgruber, P., and C. Wemmer.  2004.  National elephant symposium and workshop. Report to the USFWS and the Myanmar Forest Department.
7 Pfaff, G. 1930.  Reports on Diseases of Elephants.  Government Printing. Rangoon. 91
8 Wemmer, C., P. Leimgruber and D. S. Kelly.  2005.  Managing wild elephants in Alaungdaw Kathapa National Park and Htamanthi Wildlife Sanctuary.  Report to the USFWS and the Myanmar Forest Department.

Ganswindt, A., Rasmussen, H.B., Heistermann, M., Hodges, J.K., 2005. The sexually active states of free-ranging male African elephants (Loxodonta africana): defining musth and non-musth using endocrinology, physical signals, and behavior
652. Horm. Behav. 47, 83-91.
Abstract: Musth in male African elephants, Loxodonta africana, is associated with increased aggressive behavior, continuous discharge of urine, copious secretions from the swollen temporal glands, and elevated androgen levels. During musth, bulls actively seek out and are preferred by estrous females although sexual activity is not restricted to the musth condition. The present study combines recently established methods of fecal hormone analysis with long-term observations on male-female associations as well as the presence and intensity of physical signals to provide a more detailed picture about the physical, physiological, and behavioral characteristics of different states of sexual activity in free-ranging African elephants. Based on quantitative shifts in individual bull association patterns, the presence of different physical signals, and significant differences in androgen levels, a total of three potential sub-categories for sexually active bulls could be established. The results demonstrate that elevations in androgen levels are only observed in sexually active animals showing temporal gland secretion and/or urine dribbling, but are not related to the age of the individual. Further, none of the sexually active states showed elevated glucocorticoid output indicating that musth does not represent an HPA-mediated stress condition. On the basis of these results, we suggest that the term "musth" should be exclusively used for the competitive state in sexually active male elephants and that the presence of urine dribbling should be the physical signal used for defining this state

Ganswindt, A., Rasmssen, H.B., Heistermann, M., Hodges, J.K., 2005. The sexually active states of free-ranging male African elephants (Loxodonta africana): defining musth and non-musth using endocrinology, physical signals, and behavior. Horm Behav 47, 83-91.
Abstract: Musth in male African elephants, Loxodonta africana, is associated with increased aggressive behavior, continuous discharge of urine, copious secretions from the swollen temporal glands, and elevated androgen levels. During musth, bulls actively seek out and are preferred by estrous females although sexual activity is not restricted to the musth condition. The present study combines recently established methods of fecal hormone analysis with long-term observations on male-female associations as well as the presence and intensity of physical signals to provide a more detailed picture about the physical, physiological, and behavioral characteristics of different states of sexual activity in free-ranging African elephants. Based on quantitative shifts in individual bull association patterns, the presence of different physical signals, and significant differences in androgen levels, a total of three potential sub-categories for sexually active bulls could be established. The results demonstrate that elevations in androgen levels are only observed in sexually active animals showing temporal gland secretion and/or urine dribbling, but are not related to the age of the individual. Further, none of the sexually active states showed elevated glucocorticoid output indicating that musth does not represent an HPA-mediated stress condition. On the basis of these results, we suggest that the term "musth" should be exclusively used for the competitive state in sexually active male elephants and that the presence of urine dribbling should be the physical signal used for defining this state.

Garstang, M., 2005. Long-distance, low-frequency elephant communication. J Comp Physiol A Neuroethol Sens Neural Behav Physiol. 191, 299.
Abstract: Erratum: J Comp Physiol A Neuroethol Sens Neural Behav Physiol. 2004; Oct;190(10):791-805. Epub 2004 Sep 2. The production, transmission, and reception of and the behavioral response to long-distance, low-frequency sound by elephants is reviewed. The structure of low-frequency calls generated by elephants is separated into the "source" and the "filter" roles played by the lungs, larynx and vocal track, the composition of the expired air and the ambient air temperature. Implications regarding the size, age, sex, sexual and physical status follow from the call structure and detection. Reception of the signal is discussed in terms of the characteristics of the elephant's ear with particular attention to the determination of the threshold of hearing and the ability to locate the source of low-frequency sounds. Factors which influence the transmission of near infrasound are related to atmospheric structure. The critical role played by the thermal stratification and vertical gradient and magnitude of the wind in determining both the range and the detection of a signal are discussed for open and closed elephant habitats. Infrasound plays a pervasive role in reproduction, resource utilization, avoidance of predation and other social interactions. Current and future technology can be expected to contribute to the detection and interpretation of elephant communication. This will aid in the understanding of behavior and in efforts to sustain the species.

Glickman, S.E., Short, R.V., Renfree, M.B., 2005. Sexual differentiation in three unconventional mammals: Spotted hyenas, elephants and tammar wallabies. Hormones and Behaviour 48, 403-417.
Abstract: The present review explores sexual differentiation in three non-conventional species: the spotted hyena, the elephant and the tammar wallaby, selected because of the natural challenges they present for contemporary understanding of sexual differentiation. According to the prevailing view of mammalian sexual differentiation, originally proposed by Alfred Jost, secretion of androgen and anti-Mullerian hormone (AMH) by the fetal testes during critical stages of development accounts for the full range of sexually dimorphic urogenital traits observed at birth. Jost's concept was subsequently expanded to encompass sexual differentiation of the brain and behavior. Although the central focus of this review involves urogenital development, we assume that the novel mechanisms described in this article have potentially significant implications for sexual differentiation of brain and behavior, a transposition with precedent in the history of this field. Contrary to the ''specific'' requirements of Jost's formulation, female spotted hyenas and elephants initially develop male-type external genitalia prior to gonadal differentiation. In addition, the administration of anti-androgens to pregnant female spotted hyenas does not prevent the formation of a scrotum, pseudoscrotum, penis or penile clitoris in the offspring of treated females, although it is not yet clear whether the creation of masculine genitalia involves other steroids or whether there is a genetic mechanism bypassing a hormonal mediator. Wallabies, where sexual differentiation occurs in the pouch after birth, provide the most conclusive evidence for direct genetic control of sexual dimorphism, with the scrotum developing only in males and the pouch and mammary glands only in females, before differentiation of the gonads. The development of the pouch and mammary gland in females and the scrotum in males is controlled by genes on the X chromosome. In keeping with the ''expanded'' version of Jost's formulation, secretion of androgens by the fetal testes provides the best current account of a broad array of sex differences in reproductive morphology and endocrinology of the spotted hyena, and androgens are essential for development of the prostate and penis of the wallaby. But the essential circulating androgen in the male wallaby is 5α androstanediol, locally converted in target tissues to DHT, while in the pregnant female hyena, androstenedione, secreted by the maternal ovary, is converted by the placenta to testosterone (and estradiol) and transferred to the developing fetus. Testicular testosterone certainly seems to be responsible for the behavioral phenomenon of musth in male elephants. Both spotted hyenas and elephants display matrilineal social organization, and, in both species, female genital morphology requires feminine cooperation for successful copulation. We conclude that not all aspects of sexual differentiation have been delegated to testicular hormones in these mammals. In addition, we suggest that research on urogenital development in these non-traditional species directs attention to processes that may well be operating during the sexual differentiation of morphology and behavior in more common laboratory mammals, albeit in less dramatic fashion.

Glickman, S.E., Short, R.V., Renfree, M.B., 2005. Sexual differentiation in three unconventional mammals: spotted hyenas, elephants and tammar wallabies
566. Horm. Behav. 48, 403-417.
Abstract: The present review explores sexual differentiation in three non-conventional species: the spotted hyena, the elephant and the tammar wallaby, selected because of the natural challenges they present for contemporary understanding of sexual differentiation. According to the prevailing view of mammalian sexual differentiation, originally proposed by Alfred Jost, secretion of androgen and anti-Mullerian hormone (AMH) by the fetal testes during critical stages of development accounts for the full range of sexually dimorphic urogenital traits observed at birth. Jost's concept was subsequently expanded to encompass sexual differentiation of the brain and behavior. Although the central focus of this review involves urogenital development, we assume that the novel mechanisms described in this article have potentially significant implications for sexual differentiation of brain and behavior, a transposition with precedent in the history of this field. Contrary to the "specific" requirements of Jost's formulation, female spotted hyenas and elephants initially develop male-type external genitalia prior to gonadal differentiation. In addition, the administration of anti-androgens to pregnant female spotted hyenas does not prevent the formation of a scrotum, pseudoscrotum, penis or penile clitoris in the offspring of treated females, although it is not yet clear whether the creation of masculine genitalia involves other steroids or whether there is a genetic mechanism bypassing a hormonal mediator. Wallabies, where sexual differentiation occurs in the pouch after birth, provide the most conclusive evidence for direct genetic control of sexual dimorphism, with the scrotum developing only in males and the pouch and mammary glands only in females, before differentiation of the gonads. The development of the pouch and mammary gland in females and the scrotum in males is controlled by genes on the X chromosome. In keeping with the "expanded" version of Jost's formulation, secretion of androgens by the fetal testes provides the best current account of a broad array of sex differences in reproductive morphology and endocrinology of the spotted hyena, and androgens are essential for development of the prostate and penis of the wallaby. But the essential circulating androgen in the male wallaby is 5alpha androstanediol, locally converted in target tissues to DHT, while in the pregnant female hyena, androstenedione, secreted by the maternal ovary, is converted by the placenta to testosterone (and estradiol) and transferred to the developing fetus. Testicular testosterone certainly seems to be responsible for the behavioral phenomenon of musth in male elephants. Both spotted hyenas and elephants display matrilineal social organization, and, in both species, female genital morphology requires feminine cooperation for successful copulation. We conclude that not all aspects of sexual differentiation have been delegated to testicular hormones in these mammals. In addition, we suggest that research on urogenital development in these non-traditional species directs attention to processes that may well be operating during the sexual differentiation of morphology and behavior in more common laboratory mammals, albeit in less dramatic fashion

Leong, K.M., Burks, K., Rizkalla, C.E., Savage, A., 2005. Effects of reproductive and social context on vocal communication in captive female African elephants (Loxodonta africana). Zoo Biology 24, 331-347.
Abstract: Female African elephants advertise changes in reproductive condition to males through a variety of modalities, including an increase in low-frequency vocalizations, presumed to travel long distances.  Although males respond to these vocalizations, it has been suggested that their proximate function may be to signal to nearby females rather than to distant males. Because elephants live in a female-bonded society, it is likely  that  changes in female reproductive condition also affect close-range  interactions between high- and low-ranking females and that  vocalizations  may mediate these interactions. To examine female-female interactions related to vocal production and the ovulatory cycle, this year-long study monitored behavior, vocalizations and hormonal cycles for a group of six female captive African elephants at Disney's Animal Kingdom. Rates of several types of close-range interactions were observed to change over the phases of the estrous cycle, and rank seemed to affect whether or not low-frequency vocalizations were given in association with these interactions. Results of this study suggest that a female African  elephant's  immediate social context and rank in the social hierarchy interact with  the  hormonal cycle in the production of low-frequency vocalizations, thus  many  of these vocalizations may not function proximately as signals to  distant  males, but may be a result of the changing dynamics among females.

Nissani, M., Hoefler-Nissani, D., Lay, U.T., Htun, U.W., 2005. Simultaneous visual discrimination in Asian elephants
631. J. Exp. Anal. Behav. 83, 15-29.
Abstract: Two experiments explored the behavior of 20 Asian elephants (Elephas aximus) in simultaneous visual discrimination tasks. In Experiment 1, 7 Burmese logging elephants acquired a white+/black- discrimination, reaching criterion in a mean of 2.6 sessions and 117 discrete trials, whereas 4 elephants acquired a black+/white- discrimination in 5.3 sessions and 293 trials. One elephant failed to reach criterion in the white+/black- task in 9 sessions and 549 trials, and 2 elephants failed to reach criterion in the black+/white- task in 9 sessions and 452 trials. In Experiment 2, 3 elephants learned a large/small transposition problem, reaching criterion within a mean of 1.7 sessions and 58 trials. Four elephants failed to reach criterion in 4.8 sessions and 193 trials. Data from both the black/white and large/small discriminations showed a surprising age effect, suggesting that elephants beyond the age of 20 to 30 years either may be unable to acquire these visual discriminations or may require an inordinate number of trials to do so. Overall, our results cannot be readily reconciled with the widespread view that elephants possess exceptional intelligence

Ortolani, A., Leong, K., Graham, L., Savage, A., 2005. Behavioral indices of estrus in a group of captive African elephants (Loxodonta africana). Zoo Biology 24, 311-329.
Abstract: This study investigated behavioral signals of estrus by systematically monitoring the interactions of one male with four female African elephants housed in a naturalistic outdoor enclosure at Disney's Animal Kingdom over a period of 11 months. We measured changes in five spatial behaviors and 22 tactile-contact behaviors, as well as changes in serum progestagen and LH concentrations, across three ovarian cycles for each female. Two females did not cycle during the study. Three different phases of the ovarian cycle were identified: mid luteal, anovulatory follicular, ovulatory follicular.  The male followed more and carried out more genital inspections, flehmen, and trunk-to-mouth behaviors toward cycling females during their ovulatory phase. Genital inspections by the male peaked above baseline levels on  the  day of an LH surge, and up to 9 days before, in both cycling females  and,  thus, might be a useful behavioral index of estrus. The male also carried out more genital inspections, flehmen, and trunk touches to the back leg toward ovulatory cycling than noncycling females. Overall, our results  indicated that: 1) a single subadult African elephant male could  discriminate two females in the ovulatory phase of their cycle (i.e.,  during  the 3 weeks preceding ovulation) from the mid luteal phase; 2) the male  also  discriminated two cycling females in the ovulatory and anovulatory  follicular phases from two noncycling females; 3) two females in the  ovulatory phase of the cycle displayed a greater variety of  tactile-contact  behavior toward the male compared to the other cycle phases.

Perez-Barberia, F.J., Gordon, I.J., 2005. Gregariousness increases brain size in ungulates
586. Oecologia. 145, 41-52.
Abstract: The brain's main function is to organise the physiological and behavioural responses to environmental and social challenges in order to keep the organism alive. Here, we studied the effects that gregariousness (as a measurement of sociality), dietary habits, gestation length and sex have on brain size of extant ungulates. The analysis controlled for the effects of phylogeny and for random variability implicit in the data set. We tested the following groups of hypotheses: (1) Social brain hypothesis-gregarious species are more likely to have larger brains than non-gregarious species because the former are subjected to demanding and complex social interactions; (2) Ecological hypothesis-dietary habits impose challenging cognitive tasks associated with finding and manipulating food (foraging strategy); (3) Developmental hypotheses (a) energy strategy: selection for larger brains operates, primarily, on maternal metabolic turnover (i.e. gestation length) in relation to food quality because the majority of the brain's growth takes place in utero, and finally (b) sex hypothesis: females are expected to have larger brains than males, relative to body size, because of the differential growth rates of the soma and brain between the sexes. We found that, after adjusting for body mass, gregariousness and gestation length explained most of the variation in brain mass across the ungulate species studied. Larger species had larger brains; gregarious species and those with longer gestation lengths, relative to body mass, had larger brains than non-gregarious species and those with shorter gestation lengths. The effect of diet was negligible and subrogated by gestation length, and sex had no significant effect on brain size. The ultimate cause that could have triggered the co-evolution between gestation length and brain size remains unclear

Poole, J.H., Tyack, P.L., Stoeger-Horwath, A.S., Watwood, S., 2005. Animal behaviour: elephants are capable of vocal learning
623. Nature 434, 455-456.
Abstract: There are a few mammalian species that can modify their vocalizations in response to auditory experience--for example, some marine mammals use vocal imitation for reproductive advertisement, as birds sometimes do. Here we describe two examples of vocal imitation by African savannah elephants, Loxodonta africana, a terrestrial mammal that lives in a complex fission-fusion society. Our findings favour a role for vocal imitation that has already been proposed for primates, birds, bats and marine mammals: it is a useful form of acoustic communication that helps to maintain individual-specific bonds within changing social groupings

Roca, A.L., O'Brien, S.J., 2005. Genomic inferences from Afrotheria and the evolution of elephants
558. Curr. Opin. Genet. Dev. 15, 652-659.
Abstract: Recent genetic studies have established that African forest and savanna elephants are distinct species with dissociated cytonuclear genomic patterns, and have identified Asian elephants from Borneo and Sumatra as conservation priorities. Representative of Afrotheria, a superordinal clade encompassing six eutherian orders, the African savanna elephant was among the first mammals chosen for whole-genome sequencing to provide a comparative understanding of the human genome. Elephants have large and complex brains and display advanced levels of social structure, communication, learning and intelligence. The elephant genome sequence might prove useful for comparative genomic studies of these advanced traits, which have appeared independently in only three mammalian orders: primates, cetaceans and proboscideans

Roth, G., Dicke, U., 2005. Evolution of the brain and intelligence
612. Trends Cogn Sci. 9, 250-257.
Abstract: Intelligence has evolved many times independently among vertebrates. Primates, elephants and cetaceans are assumed to be more intelligent than 'lower' mammals, the great apes and humans more than monkeys, and humans more than the great apes. Brain properties assumed to be relevant for intelligence are the (absolute or relative) size of the brain, cortex, prefrontal cortex and degree of encephalization. However, factors that correlate better with intelligence are the number of cortical neurons and conduction velocity, as the basis for information-processing capacity. Humans have more cortical neurons than other mammals, although only marginally more than whales and elephants. The outstanding intelligence of humans appears to result from a combination and enhancement of properties found in non-human primates, such as theory of mind, imitation and language, rather than from 'unique' properties

Soltis, J., Leong, K., Savage, A., 2005. African elephant vocal communication II: Rumble variation reflects the individual identity and emotional state of callers. Animal Behaviour 70, 589-599.
Abstract: The most common vocalization of the African elephant, Loxodonta  africana, is the rumble, but there is no consensus as to how many rumble subtypes exist. From the standpoint of social function, many types of rumble have been proposed. From a structural standpoint, however, few studies have examined detailed acoustic measurements of a large number of calls. We analysed 270 rumbles from six adult female African elephants housed at Disney's Animal Kingdom (Lake Buena Vista, Florida, U.S.A.). Subjects wore collars outfitted with microphones and radiotransmitters that allowed recording of vocalizations from identified individuals. Rumble vocalizations were digitized and both source and filter features were measured for each call. Behavioural and endocrine data were collected so that acoustical data could be placed into the context of ongoing social behaviour and reproductive state. Multidimensional scaling analysis revealed that, from a structural standpoint, rumbles from this captive setting could not be divided into distinct subtypes, but there was extensive acoustic variation across rumbles. Discriminant function analysis and MANOVA were employed to further explore this variation. First, acoustic characteristics varied according to the individual identity of the caller. Second, rumbles varied as a function of negative emotional arousal. When associating with dominant animals, subordinate females produced rumbles with lower cepstral coefficients, suggesting low tonality and unstable pitch in the voice, compared to rumbles produced outside of the presence of dominant animals. Rumbles as a whole did not cluster into distinct acoustic types, but structural variation in rumbles reflected the individual identity and emotional state of callers.

Soltis, J., Leong, K., Savage, A., 2005. African elephant vocal communication I: Antiphonal calling behaviour among affiliated females. Animal Behaviour 70, 579-587.
Abstract: African elephants, Loxodonta africana, are well known for their use of a low-frequency 'rumble' vocalization, which is thought to function in long-distance communication. Less work, however, has been conducted on short-distance communication within groups, and on spontaneously occurring vocal exchanges among identified individuals in particular. This is due in part to the fact that low-frequency rumbles are difficult to assign to individual callers. We collected vocal data on a group of six female African elephants housed at Disney's Animal Kingdom to determine whether they exchange rumbles in alternating sequences (also known as antiphonal calling). Subjects wore collars outfitted with microphones and radiotransmitters that allowed identification of individual callers, and behavioural and endocrine data were collected so that vocal activity could be examined in the context of social behaviour and reproductive state. First, we found that females did not produce rumbles at random, but were nearly twice as likely to produce rumbles shortly after rumbles from other group members. Second, the relative dominance rank and reproductive state of callers did not affect the probability of vocal response, but affiliative relationship with the caller had a strong influence on rumble response.  Females were most likely to respond in kind to the rumbles of their most affiliated partners compared to less affiliated group members. Third, video analysis showed that rumble exchanges occurred in variable contexts, including when animals were out of contact, during reunions, and while in close proximity. Also, affiliated partners often vocalized in sequence when approached by dominant individuals. The results of these analyses show that affiliated female African elephants exchange rumbles antiphonally, and imply multiple functions for such vocal exchanges.

Vidya, T.N.C., Sukumar, K., 2005. Social organization of the Asian elephant (Elephas maximus) in southern India inferred from microsatellite DNA. J Ethol 23, 205-210.
Abstract: Social organization of the Asian elephant (Elephas maximus) is not well understood in the absence of long-term studies of identified individuals. Adult Asian elephant females and their young offspring of both sexes form matriarchal groups, with pubertal males dispersing from natal groups, but whether these social groups represent families and whether males show locational or social dispersal were unknown. Using nuclear microsatellite loci amplified from dung-extracted DNA of free-ranging elephants in a large southern Indian population, we demonstrate that female-led herds comprise closely related individuals that are indeed families, and that males exhibit non-random locational dispersal.

Wittemyer, G., Douglas-Hamilton, I., Getz, W.M., 2005. The socioecology of elephants: Analysis of the processes creating multitiered social structures. Animal Behaviour.
Abstract: In this paper, we investigate the formation and function of the multilevelled, fission-fusion social structure in a free-ranging African elephant, Loxodonta africana, population. We quantitatively identified the existence of four social tiers by using cluster analysis on individual association data. We assessed the effects of season and study period on social structuring and levels of cohesion within and among social units. We found that second-tier units, potentially the equivalent of the 'family', were stable across seasonal periods but the number of units increased as the study progressed and the population grew. It appears that these units were sufficiently small not to be influenced by ecologically related factors, such as resource competition, that might otherwise lead to them splitting. On the other hand, third- and fourth tier units were significantly affected by season in a way that suggests a trade-off between ecological costs (e.g. from resource competition) and different social and ecological benefits (e.g. from predator defence, territoriality, knowledge sharing and rearing of young). Age structure also appeared to influence this multitiered social organization. The size of second-tier social units was significantly affected by the age of matriarchs: units lead by matriarchs likely to be grandmothers (i.e. females 35 years and older) were significantly larger than those lead by younger matriarchs. We present a conceptual framework for understanding the emergence of multiple-tier social structure from interactions driven by socioecological processes. This study is the first to use rigorous quantitative methods to statistically show the existence of four hierarchical tiers of social organization in a nonhuman animal. Additionally, our results elucidate the role that ecological processes play in producing complex social structures.

 2004. Elephant Husbandry Resource Guide. International Elephant Foundation, Azle. TX.

Alter, S., 2004. Elephas maximus: A portrait of the Indian elephant. Harcourt Press.
Abstract: Review from Amazon: Mixing mythology and natural history, Stephen Alter lets readers share his lifelong love for the Indian elephant, Elephas maximus. While legends threaten to overwhelm facts in the tale, Alter has nonetheless presented an accurate portrait of his subject, true to centuries of Indian tradition. Beyond metaphors and fables, elephants occupy an important place in Sanskrit literature. Gajashastra, or "elephant science," was studied and recorded in several texts that are based on oral traditions.As much travelogue as science book, Elephas inextricably links the Indian elephant with the history of southern Asia itself. In pre-colonial India, elephants were wound up in religion and daily life; in modern times, the animals were first hunted then fetishized by Westerners. Alter reserves judgment on these issues, except to note that none of India's 20th-century history has been good for elephant populations, which are endangered or threatened nearly everywhere. He treks into parks and reserves, seeking out wild elephants and describing their awe-inspiring behaviors. The stories he uncovers along the way--of temple elephants, mysterious Elephanta  Island, seagoing elephants, and the god Ganesha--weave a spellbinding tale. --Therese Littleton

Bradshaw, I.G.A., 2004. Not by Bread Alone: Symbolic Loss, Trauma, and Recovery in Elephant Communities. Society And Animals 12, 143-158.
Abstract: Like many humans in the wake of genocide and war, most wildlife today has sustained trauma. High rates of mortality, habitat destruction, and social breakdown precipitated by human actions are unprecedented in history. Elephants are one of many species dramatically affected by violence. Although elephant communities have processes, rituals, and social structures for responding to trauma - grieving, mourning, and socialization - the scale, nature, and magnitude of human violence have disrupted their ability to use these practices. Absent the cultural, carrier groups (murdered elephant matriarchs and elders) who traditionally lead and teach these healing practices, humans must assume the role. Trauma theory has brought attention to victims' severe, sustained psychological damage. Looking through the lens of trauma theory provides a better understanding of how systematic violence has affected individuals and groups and how the pervasive nature of traumatic events affects human-nonhuman animal relationships. The framing of recent trauma theory compels conservationists to create new relationships - neither anthropocentric nor powerbased - with nonhuman animals. The David Sheldrick Wildlife Trust, Kenya, shows how humans, taking on the role of interspecies witness, bring orphan elephants back to health and help re-build elephant communities shattered by genocide.

Burks, K.D., Mellen, J.D., Miller, G.W., Lehnhardt, J., Weiss, A., Figueredo, A.J., Maple, T.L., 2004. Comparison of two introduction methods for African elephants (Loxodonta africana). Zoo Biology 23,  109-126.
Abstract:
Managers must consider an animal's potential for aggression when they decide to change or form a captive social group formation. In this study we compared two introduction methods (termed "sequential" and "nonsequential" introductions) in African elephants to assess their effectiveness in managing aggression and minimizing stress. Both introduction methods included four phases: baseline, visual contact, limited tactile contact, and physical introduction. In the sequential introduction, these steps were followed sequentially, and empirical data were considered during decision-making. In the nonsequential introduction, these steps were not followed sequentially, and decision-making was based primarily on intuitive assessments by animal managers. Behavioral data and fecal corticoid concentrations were measured throughout both types of introduction. The behavior categories measured included active aggression, passive aggression, submissive behavior, undesirable/stress-related behavior, and affiliative behavior. While the role of affiliative behavior was surprising, general behavior patterns were characterized by increases in behavior as animals progressed to the next phase of introduction regardless of introduction type. These increases then attenuated over time during each phase. Overall, less behavior was observed during the sequential introduction, as predicted.  The data suggest that the sequential introduction managed aggression more effectively. Similar patterns were predicted for undesirable/stress-related behavior and fecal corticoid concentration. Undesirable/stress-related behavior was a poor predictor of observed behavior patterns. Although the patterns differed from those predicted, higher concentrations of fecal corticoids were measured during the nonsequential introduction and correlated significantly only with submissive behavior. While more investigation is warranted, the data indicate that the nonsequential introduction brought about an increased physiological response. Overall, the sequential introduction method appeared to manage aggression and stress better than the nonsequential technique. Every introduction is subject to factors that can influence success, such as staff experience, the design of the facility, and the animals' social histories. It is hoped that the rigorous sequential protocol will be a useful tool in the animal manager's "toolbox" for planning and implementing introductions. Applications of this introduction method are also discussed.

Cerling, T.E., Passey, B.H., Ayliffe, L.K., Cook, C.S., Ehleringer, J.R., Harris, J.M., Dhidha, M.B., Kasiki, S.M., 2004. Orphans' tales: seasonal dietary changes in elephants from Tsavo National Park, Kenya. Palaeogeography Palaeoclimatology Palaeoecology 206, 367-376.
Abstract: The similarity of delta(13)C and delta(15)N patterns in hairs of different individuals from the Tsavo East orphaned elephant herd indicates that a single hair represents the dietary preferences and behavior of the entire group. Multiple tail hairs from the same individual collected at different times allows a chronology to be established because of the overlap in isotope patterns in hair, and there is a very high correlation between
hair from different individuals in the same group. Forward modeling using a three-component isotope turnover model for hair allows a precise estimate of diet of these elephants over a 2-year interval. Elephants from Tsavo East National Park in Kenya feed predominantly on C-3 leaves, although they have a significant fraction Of C-4 grass in their diet for a short time at the beginning of the rainy season. The overall integrated diet for the elephants studied is between 10% and 15% C-4 grass, although it reaches up to 60% for short intervals. Stable carbon isotope analyses of elephant tooth enamel show that the average integrated dietary preference of elephants in Tsavo National Park remained less than 25% grass between 1940 and the present.

de Oliveira, C.A., West, G.D., Houck, R., Leblanc, M., 2004. Control of musth in an Asian elephant bull (Elephas maximus) using leuprolide acetate
716. J. Zoo. Wildl. Med. 35, 70-76.
Abstract: The results of long-term administration of leuprolide acetate (LA) depot in a 52-yr-old Asian elephant bull (Elephas maximus) for control of musth are presented. Twelve injections were administered for 6 yr during our interpretation of early musth or "premusth." Intervals between musth periods during the study varied from 2 to 34 mo. Blood samples, drawn weekly, were assayed for serum testosterone concentrations; mean levels were 11.78 +/- 1.97 nmol/L throughout the first 26 mo of the study, 7.28 +/- 1.28 nmol/L during the following 21 mo, and 0.45 +/- 0.035 nmol/L in the last 34 mo of this study. Early musth signs ceased within 3 days of drug administration after 10 of 12 injections. The mean serum testosterone concentrations were significantly decreased by the last 34 mo of the study. The results suggest leuprolide is a suitable alternative for controlling or preventing (or both) musth in captive Asian elephants, although permanent reproductive effects may occur. Zoos and wildlife conservation institutions could benefit from the use of LA in Asian elephants to increase the male availability in captivity, consequently ensuring genetic diversity and the perpetuation of the species

Dembiec, D.P., Snider, R.J., Zanella, A.J., 2004. The effects of transport stress on tiger physiology and behavior. Zoo Biology 23, 335-346.
Abstract: Tigers are often transported for education, conservation, and zoo enhancement purposes, however the effect of transfer on them has not yet been documented. Our objective was to evaluate how transport affects the behavior and physiology of tigers, taking into account previous experience with the transport procedure. We simulated transport by relocating naive tigers in a small individual transfer cage. Two tigers had prior experience with the procedure, and three tigers were naive to it. After 30 min, each tiger was released back into their original enclosure. Physiological measurements were recorded for four of the naive tigers; these included respiration rate and immune-reactive fecal cortisol response using radioimmunoassay. We also recorded the behavior of all naive tigers before, during, and after transport. Our behavioral analysis included activity level, pacing behavior, time spent investigating, respiration rate, and ear position. Average respiration rates of all tigers increased from 56.1 breaths/min to 94.6 breaths/min during transport and to 132.3 breaths/min 10 min following release into their enclosures. Average immune-reactive cortisol concentrations peaked 3-6 days after transport at 239% above baseline and returned to baseline levels 9-12 days afterward. During their peak time block, naýve tigers exhibited a higher average increase in cortisol levels (482% above baseline) than the experienced tigers (158% above baseline). The naýve tigers' average immune-reactive cortisol concentration remained elevated for a longer period (9-12 days) than the experienced tigers' (3-6 days). In both groups, behavioral responses ranged from active to inactive, however naýve tigers performed these repertoires with greater intensity by pacing faster and performing fewer state changes. Results suggest that prior exposure to elements of the transport procedure may lead to some level of habituation, thus reducing the effects of transportation stress.

Freeman, E.W., Weiss, E., Brown, J.L., 2004. Examination of the interrelationships of behavior, dominance status, and ovarian activity in and African elephants. Zoo Biology 23, 431-448.
Abstract: Ovarian inactivity has been identified in captive African (Loxodonta africana) and Asian (Elephas maximus) elephants and is thought to be mediated in part by social influences. Thus, a survey was conducted to determine how behavior and dominance status relate to each other and to ovarian cyclicity. For both Asian and African elephants, dominance status was positively correlated with relative size, age, temperament, disciplinary nature, and willingness to share novel objects. Relative size and temperament were also related to disciplinary nature toward herdmates. Behavior toward keepers was a good indicator of the willingness of elephants to follow cmmands, whereas sharing novel objects was positively correlated with object curiosity. Finally, dominance status, temperament, and disciplinary nature were all correlated with willingness to share. Comparisons of ovarian cyclicity status with behavior rankings were conducted only for African elephants because of the low number of noncycling Asian elephants surveyed. Overall, social status appeared to be the best predictor of ovarian activity in African elephants. Noncycling African elephant females ranked higher in the dominance hierarchy and gave more discipline to herdmates than cycling cohorts. It remains to be determined whether these are cause or effect relationships, but clearly it is important to understand how physical and social attributes impact physiological processes, such as reproduction. Captive management now needs to focus on optimizing social and environmental conditions to maximize reproductive potential in elephants.

Garstang, M., 2004. Long-distance, low-frequency elephant communication. J Comp Physiol A 190, 791-805.
Abstract: The production, transmission, and reception of and the behavioral response to long-distance, low-frequency sound by elephants is reviewed. The structure of low-frequency calls generated by elephants is separated into the ''source'' and the ''filter'' roles played by the lungs, larynx and vocal track, the composition of the expired air and the ambient air temperature. Implications regarding the size, age, sex, sexual and physical status follow from the call structure and detection. Reception of the signal is discussed in terms of the characteristics of the elephant's ear with particular attention to the determination of the threshold of hearing and the ability to locate the source of low-frequency sounds. Factors which influence the transmission of near infrasound are related to atmospheric structure. The critical role played by the thermal stratification and vertical gradient and magnitude of the wind in determining both the range and the detection of a signal are discussed for open and closed elephant habitats. Infrasound plays a pervasive role in reproduction, resource utilization, avoidance of predation and other social interactions. Current and future technology can be expected to contribute to the detection and interpretation of elephant communication. This will aid in the understanding of behavior and in efforts to sustain the species.

Garstang, M., 2004. Long-distance, low-frequency elephant communication
686. J. Comp Physiol A Neuroethol. Sens. Neural Behav. Physiol 190, 791-805.
Abstract: The production, transmission, and reception of and the behavioral response to long-distance, low-frequency sound by elephants is reviewed. The structure of low-frequency calls generated by elephants is separated into the "source" and the "filter" roles played by the lungs, larynx and vocal track, the composition of the expired air and the ambient air temperature. Implications regarding the size, age, sex, sexual and physical status follow from the call structure and detection. Reception of the signal is discussed in terms of the characteristics of the elephant's ear with particular attention to the determination of the threshold of hearing and the ability to locate the source of low-frequency sounds. Factors which influence the transmission of near infrasound are related to atmospheric structure. The critical role played by the thermal stratification and vertical gradient and magnitude of the wind in determining both the range and the detection of a signal are discussed for open and closed elephant habitats. Infrasound plays a pervasive role in reproduction, resource utilization, avoidance of predation and other social interactions. Current and future technology can be expected to contribute to the detection and interpretation of elephant communication. This will aid in the understanding of behavior and in efforts to sustain the species

Lazar, J., Rasmussen, L.E., Greenwood, D.R., Bang, I.S., Prestwich, G.D., 2004. Elephant albumin: a multipurpose pheromone shuttle
691. Chem. Biol. 11, 1093-1100.
Abstract: (Z)-7-dodecenyl acetate (Z7-12:Ac) is present in the urine of female Asian elephants (Elephas maximus) approaching ovulation and functions as a female-to-male sex pheromone. Here we show that a significant fraction of the pheromone in the urine is bound to a protein, elephant serum albumin (ESA), and provide evidence for key physiological functions of urinary ESA. Our biochemical and behavioral experiments suggest a three-fold role of ESA in pheromone signaling: (1) transporting Z7-12:Ac from serum into urine; (2) extending the presence of the pheromone in the environment without hampering detection; and (3) targeting pheromone delivery to chemosensory organs through localized release of the ligand induced by a pH change. The exploitation of albumin in pheromone transport clearly distinguishes the elephant from other mammals studied, and complements the uniqueness of elephant anatomy, physiology, and behavior

Rees, P.A., 2004. Low environmental temperature causes an increase in stereotypic behaviour in captive Asian elephants (Elephas maximus). Journal of Thermal Biology 29, 37-43.
Abstract: (1) Captive Asian elephants exhibited a strong negative correlation between  maximum daily temperature and the frequency of stereotypic behaviour, in those animals predisposed to exhibit stereotypies. (2) The frequency of stereotypic behaviours increased throughout each study day (1000-1400 h), reaching maximum values at the end of the day, as feeding time approached. (3) The frequencies of stereotypic behaviour were higher at all times of the day on the 10 coldest days (mean maximum daily temperature=9.0degreesC) than on the 10 warmest days (mean maximum daily temperature=23.2degreesC) of the study. (4) There was a strong negative correlation between body mass and mean frequency of stereotypic behaviour. This may have been because smaller animals lost heat faster than larger animals. (5) The aetiology of stereotypic behaviour was unclear, but it was likely to be the result of poor husbandry experienced in early life, such as chaining and inappropriate housing. Hunger and the physical thwarting of attempts to reach food and shelter may have been the proximate cause of individual episodes of stereotypic behaviour, with temperature acting as a compounding factor. There was no evidence that stereotypic behaviour developed in response to exposure to cold.

Rees, P.A., 2004. Some preliminary evidence of the social facilitation of mounting behavior in a juvenile bull asian elephant (Elephas maximus)
734. J. Appl. Anim Welf. Sci. 7, 49-58.
Abstract: This study recorded sexual behavior within a captive herd of 8 Asian elephants for approximately 230 hr on 50 days over a period of 10 months. The study observed a single adult and a single juvenile bull mounting cows more than 160 times. When the juvenile bull was between 4 years, 2 months and 4 years, 8 months old, he exhibited mounting behavior only on days when adult mounting occurred. Adult mounting always occurred first. Beyond the age of 4 years, 8 months, the juvenile bull exhibited spontaneous mounting behavior in the absence of adult mounting. This suggests that mounting behavior may develop because of social facilitation. Determining the significance of the presence of sexually active adults in the normal development of sexual behavior in juveniles will require further studies. Encouraging the establishment of larger captive herds containing adults and calves of both sexes-if their presence is important-would improve the welfare of elephants in zoos and increase their potential conservation value

Wilson, M.L., Bloomsmith, M.A., Maple, T.L., 2004. Stereotypic swaying and serum cortisol concentrations in three captive African elephants (Loxodonta africana). Animal-Welfare 13, 39-43.
Abstract: The behaviour and serum cortisol concentrations of three captive female African elephants (Loxodonta africana) were studied to determine whether their stereotypic swaying was more prevalent before regularly scheduled events in the elephants' routine, and whether the elephants that exhibited more stereotyped swaying had lower mean serum cortisol concentrations. Behavioural data were collected during hour-long observations balanced across three periods, and during 15-min observations prior to the elephants being moved to different portions of their enclosure. Observational data were collected using instantaneous focal sampling of behaviours every 30 s. Serum cortisol measures were obtained through weekly blood withdrawal from the elephants' ears. Of the three elephants, two exhibited stereotyped swaying, which accounted for a mean of 0.4% of the scans during the hour-long observations and a mean of 18% of the scans prior to the elephants being moved between different parts of the enclosure. Swaying was highly variable among the individual elephants during both categories of observations. Additionally, both elephants swayed more prior to moving in the afternoon than prior to moving in the morning. Analyses of serum cortisol concentrations indicated that each elephant had a different mean cortisol level, which did not clearly correspond with the expression of swaying. The findings indicate that a rigidly scheduled management event may elicit stereotyped swaying in the studied elephants. Future research should document the behavioural and physiological effects of an altered management routine to improve captive elephant welfare.

Bhattacharyya, B.K., 2003. Some reproductive traits of elephants. In: Das, D. (Ed.), Healthcare, Breeding and Management of Asian Elephants. Project Elephant. Govt. of India, New Delhi, pp. 37-41.

Chakraborty, A., 2003. Common words used in elephant management. In: Das, D. (Ed.), Healthcare, Breeding and Management of Asian Elephants. Project Elephant. Govt. of India, New Delhi, pp. 191-194.

Eggert, L.S., Eggert, J.A., Woodruff, D.S., 2003. Estimating population sizes for elusive animals: the forest elephants of Kakum National Park, Ghana. Mol Ecol 12, 1389-1402.
Abstract: African forest elephants are difficult to observe in the dense vegetation, and previous studies have relied upon indirect methods to estimate population sizes. Using multilocus genotyping of noninvasively collected samples, we performed a genetic survey of the forest elephant population at Kakum National Park, Ghana. We estimated population size, sex ratio and genetic variability from our data, then combined this information with field observations to divide the population into age groups. Our population size estimate was very close to that obtained using dung counts, the most commonly used indirect method of estimating the population sizes of forest elephant populations. As their habitat is fragmented by expanding human populations, management will be increasingly important to the persistence of forest elephant populations. The data that can be obtained from noninvasively collected samples will help managers plan for the conservation of this keystone species. Ecology, Behavior and Evolution Section, Division of Biological Sciences, University of California, San Diego 92093-0116, USA. lori_eggert@hotmail.com

Leong, K.M., Ortolani, A., Graham, L.H., Savage, A., 2003. The use of low-frequency vocalizations in African elephant (Loxodonta africana) reproductive strategies. Horm Behav. 43, 433-443.
Abstract: Fertility-advertisement calls in females are predicted to occur in nonmonogamous species where males and females are widely separated in space. In African elephants, low-frequency vocalizations have thus been suggested as a reproductive strategy used by fertile females to attract mates. This study examined the use of low-frequency vocalizations with respect to different phases of the estrous cycle in African elephants by simultaneously monitoring vocalizations, behavior, and hormonal profiles. Subjects were one male and six female African elephants housed at Disney's Animal Kingdom. No acoustically distinct vocalizations were restricted to the ovulatory follicular phase. However, overall rate of low-frequency vocalization as well as the rate of one acoustically distinct vocalization changed over the estrous cycle, with highest rates of calling related to the first period of follicular growth, or anovulatory follicular phase. Elevated rates of vocalization thus were not restricted to behavioral estrus and occurred much earlier in the estrous cycle than in most species that produce fertility-advertisement calls. Both herd composition and elephant identity also affected rates of vocalization. Vocalizations therefore may not be reliable signals of actual fertility. However, the increase in vocalizations in advance of estrus may attract males to the herd prior to ovulation, facilitating both male-male competition and female choice. Once present in the herd, males may then switch strategies to use more reliable chemical and visual cues to detect ovulating females. Disney's Animal Kingdom, Lake Buena Vista, FL 32830, USA. Kristen.Leong@disney.com

Martin, F., Niemitz, C., 2003. "Right-trunkers" and "left-trunkers": side preferences of trunk movements in wild Asian elephants (Elephas maximus). J. Comp Psychol. 117, 371-379.
Abstract: In this article, the side preferences of feeding-related trunk movements of free-ranging Asian elephants (Elephas maximus) were investigated for the first time. It is hypothesized that a functional asymmetry of the trunk is necessary to perform skillful feeding movements more efficiently. This might be connected with a corresponding hemispheric specialization. Video recordings of 41 wild elephants provided frequencies and durations of the following trunk-movement categories: object contact, retrieval, and reaching. In each category, individual side preferences were found. The strength of side preferences varied between the trunk-movement categories and the sexes. Mean durations of retrieval and reaching correlated negatively with the strength of side biases. Comparing the side preferences in the unpaired trunk with analogous phenomena in other unpaired grasping organs and in primate handedness. the authors discuss possible explanations for the evolution of asymmetries in unpaired grasping organs.

Mikota, S.K., Hammatt, H., Finnegan, M. Occurrence and prevention of capture wounds in Sumatran elephants (Elephas maximus sumatranus). Proc Amer Assoc Zoo Vet.  291-293. 2003.
Ref Type: Conference Proceeding
Abstract: The capturing of elephants in Indonesia began in 1986 as an attempted solution to human-elephant conflict.  The intent was to train "problem" elephants for use in agriculture, logging and tourism.  The initial captures were conducted under the guidance of Thai mahouts and Thai koonkie elephants (trained elephants used for capture).  A number of the Indonesians that were originally trained in capture techniques still work for the government forestry department (KSDA).  The younger pawangs (elephant handlers) that participate in captures have learned from their peers.  There is no formal training program. The actual mortality rate associated with elephant captures in Sumatra is unknown as official reports are lacking.  The age structure of the existing ~ 400 captive elephants is young (most under 25) which suggests that smaller, younger elephants are preferentially captured and / or that adult elephants do not survive the capture and training processes.  Our personal experiences (Mikota and Hammatt) in Sumatra show that mortality in newly captured elephants is high.In 2001, with endorsement from the World Wide Fund for Nature-Indonesia (WWF), the Wildlife Conservation Society (WCS), Fauna and Flora International (FFI), and the International Elephant Foundation (IEF), we requested a two-year Moratorium on elephant captures during which time capture techniques would be improved and alternative conflict mediation techniques evaluated.
A Moratorium against placing additional elephants into the Elephant Training Centers has been issued by the central government, however capture for translocation is still sanctioned.  Unfortunately, the provincial governments have increasingly acted in their own interests since the government of Indonesia began a de-centralization process a few years ago. Riau Province is thought to have the largest remaining populations of wild Sumatran elephants.Fifty-seven, human-elephant conflicts occurred in Riau between 1997-2000.  Although Riau is a hotbed of conflict, problems are occurring throughout Sumatra and we are aware of conflicts and captures in Bengkulu and North Sumatra. In October 2002, we were invited by KSDA (the provincial forestry department) to accompany their team into the field as they attempted to capture a large bull that had been raiding a palm oil plantation.  This opportunity was invaluable as we were able to observe first hand the techniques being used and where improvements were needed.  As a result of this and other experiences with newly captured elephants we observed: ·Equipment (Palmer) is old, poorly maintained, and used improperly. · Essential supplies are lacking or homemade substitutes are used.
·The dose of xylazine is very high compared to wild elephant capture doses used in India and Malaysia.  The same dose is often used regardless of the size of the elephant. ·The needles are too short to reach muscle; open-ended needles are used which can become plugged with tissue, thus preventing injection. ·Neither the correct charge nor the correct load is selected.  We observed that many darts bounced making it difficult to ascertain the amount of drug injected or its depth of penetration.  Selection of an inappropriate charge results in unnecessary trauma. ·The preparation and use of darts, needles, and syringes lacks basic hygiene. ·Dart wounds are not treated and antibiotics are not administered.  · There is no understanding of stress or capture myopathy. ·The capture team was not aware that sternal recumbency severely compromises respiration in elephants and that they can quickly die in this position. ·It is believed that elephant restraints must inflict pain to prevent wild elephants from escaping once captured.  ·There is no veterinarian on the capture team. The current capture techniques result in leg wounds from unprotected chains, neck wounds from "kahs" (neck yokes made of wood and wire), and abscesses from inappropriately administered darts.  Leg and neck wounds often become maggot infested.  Infections from dart wounds are, however, the primary cause of capture-related mortality.  These abscesses can drain for several months, even with treatment, and often progress to a necrotizing fasciitis, acute sepsis, and death. The Riau Province KSDA Team has been receptive to suggested changes to minimize wounds. Provision of heavier chains has alleviated the fear that elephants will escape.  Covering the chains with fire hose or heavy plastic minimizes injuries to legs and use of the kah has been discontinued.  A basic dart wound treatment protocol has been established. In June 2003, a comprehensive Elephant Immobilization and Translocation Workshop for Sumatra is planned to retrain all of Sumatra's field teams and to upgrade equipment. Sumatra's wild elephant population probably numbers fewer than 3000 and is under continued threat.  With so few elephants left, the preservation of as many viable herds as possible takes on increased urgency.  The Moratorium achieved in 2001 has set the groundwork for KSDA to choose translocation of wild elephants rather than capture and placement into already over-crowded and under-resourced Elephant Training Centers.  We cannot guarantee that Sumatra will capture elephants only for translocation, and it is inevitable that many more elephants will end up in captivity.  Regardless, all of the elephants that must suffer the interruption of their lives at the hand of man deserve, at the very least, humane treatment.  Translocations are neither simple nor a complete panacea.  Identifying suitable translocation areas and insuring that elephants remain there are significant challenges.  WWF-Indonesia is continuing its efforts to secure the lowland forest of Tesso Nilo in Riau Province as a "safe haven" for at least some of Sumatra's wild elephants (see WWF AREAS Program – Riau, Sumatra: http://www.worldwildlife.org/species/attachments/riau_profile.pdf).  The identification of interim release sites, together with improved capture techniques, offers the hope that fewer elephants will be removed from the wild.   ACKNOWLEDGMENTS: Our work in Sumatra has been supported by the Guggenheim Foundation, a CEF grant from the American Zoo and Aquarium Association, the International Elephant Foundation, Oregon Zoo, Columbus Zoo, Disney, Peace River Refuge, the Elephant Managers Association, the Riddles Elephant and Wildlife Sanctuary, Tulsa Zoo, Toronto Zoo, Niabi Zoo, San Antonio Zoo, Denver Zoo (AAZK Chapter), Milwaukee Zoo (AAZK Chapter), the Audubon Nature Institute (Youth Volunteers), Buttonwood Park Zoo, Melbourne Zoo, and private donors.  Special thanks to Harry Peachey, John Lehnhardt, Holly Reed, Kay Backues, Mike Keele, Steve Osofsky, and Heidi and Scott Riddle.

Nath, K.C., 2003. Oestrous cycle and pregnancy in elephant. In: Das, D. (Ed.), Healthcare, Breeding and Management of Asian Elephants. Project Elephant. Govt. of India, New Delhi, pp. 42-44.

Payne, K.B., Thompson, M., Kramer, L., 2003. Elephant calling patterns as indicators of group size and composition: The basis for an acoustic monitoring system. African Journal of EcologyYear 41.
Abstract: The paper gives evidence that the vocal activity of elephants varies with group size, composition and reproductive status, and that elephants' Calling patterns could therefore provide the basis for a remote monitoring system. We examined a 3-week set of array-based audio recordings of savanna elephants (Loxodonta africana), searching for diagnostic acoustic parameters. An acoustic array made it possible to locate recorded sounds and attribute the calls to particular elephants or elephant groups. Simultaneous video recordings made it possible to document visible behaviour and roughly correlate it with vocalizations. We compared several measures of call density in elephant groups containing up to 59 individuals, and found that rates of calling increased with increasing numbers of elephants. We divided all call events into three structural types (single-voice low-frequency calls, multiple-voice clustered low-frequency calls, and single-voice high frequency calls), and found that the incidence of these varies predictably with group composition. These results suggest the value of a network of listening systems in remote areas for the collection of information on elephant abundance and population structure.

Rasmussen, L.E., Greenwood, D.R., 2003. Frontalin: a chemical message of musth in Asian elephants (Elephas maximus). Chemical Senses 28, 433-446.
Abstract: Musth is an important male phenomenon affecting many aspects of elephant society including reproduction. During musth, the temporal gland secretions (as well as the urine and breath) of adult male Asian elephants (Elephas maximus) discharge a variety of malodorous compounds together with the bicyclic ketal, frontalin. In contrast, teenage male elephants in musth release a sweet-smelling exudate from their facial temporal gland. We recently demonstrated that the concentration of frontalin becomes increasingly evident as male elephants mature. In the present study, we demonstrate that behaviors exhibited towards frontalin are consistent and dependent on the sex, developmental stage and physiological status of the responding conspecific individual. To examine whether frontalin functions as a chemical signal, perhaps even a pheromone, we bioassayed older and younger adult males, and luteal- and follicular-phase and pregnant females for their chemosensory and behavioral responses to frontalin. Adult males were mostly indifferent to frontalin, whereas subadult males were highly reactive, often exhibiting repulsion or avoidance. Female chemosensory responses to frontalin varied with hormonal state. Females in the luteal phase demonstrated low frequencies of responses, whereas pregnant females responded significantly more frequently, with varied types of responses including those to the palatal pits. Females in the follicular phase were the most responsive and often demonstrated mating-related behaviors subsequent to high chemosensory responses to frontalin. Our evidence strongly suggests that frontalin, a well-studied pheromone in insects, also functions as a pheromone in the Asian elephant: it exhibits all of the determinants that define a pheromone and evidently conveys some of the messages underlying the phenomenon of musth. Department of Biochemistry and Molecular Biology, OGI School of Science and Engineering, Oregon Health & Science University, Beaverton, OR 97006-8921, USA. betsr@bmb.ogi.edu

Sarma, K.K., 2003. Managing troublesome bulls with special reference to musth in captive Asian elephants. In: Das, D. (Ed.), Healthcare, Breeding and Management of Asian Elephants. Project Elephant. Govt. of India, New Delhi, pp. 58-66.

West, J.B., Fu, Z., Gaeth, A.P., Short, R.V., 2003. Fetal lung development in the elephant reflects the adaptations required for snorkeling in adult life. Respir Physiol Neurobiol 138, 325-333.
Abstract: The adult elephant is unique among mammals in that the pleural membranes are thickened and the pleural cavity is obliterated by connective tissue. It has been suggested that this peculiar anatomy developed because the animal can snorkel at depth, and this behavior subjects the microvessels in the parietal pleura to a very large transmural pressure. To investigate the development of the parietal pleura, the thickness of the endothoracic fascia (ET) was measured in four fetal African elephants of approximate gestational age 111-130 days, and the appearances were compared with those in human, rabbit, rat and mouse fetuses of approximately the same stage of lung organogenesis. The mean thicknesses of ET in the elephant, human, rabbit, rat and mouse were 403, 53, 29, 27 and 37 microm, respectively. This very early development of a thick parietal pleura in the elephant fetus is consistent with the hypothesis of a long history of snorkeling in the elephant's putative aquatic ancestors. Department of Medicine, University of California San Diego, 9500 Gilman Drive, La Jolla, CA 92093-0623, USA. jwest@ucsd.edu

Whitehouse, A.M., Schoeman, D.S., 2003. Ranging behaviour of elephants within a small, fenced area in Addo Elephant National Park, South Africa. African Zoology 38, 95-108.
Abstract: The elephant population (n = 324, December 2000) in Addo Elephant National Park (AENP), South Africa, is restricted to an area (103 km(2)) considerably smaller than most elephant ranges. The Addo elephants' ranging behaviour was studied in order to determine whether natural patterns of male and female ranging behaviour can be maintained within a confined area. Radio-tracking was used to facilitate measurements of hourly distances travelled, larger scale directional movements, home range sizes, centres of activity and associations. Female home ranges overlap, and interactions between family groups are frequently observed. Females and non-musth males travel similar distances and speeds on an hourly basis, but female core range sizes are larger, resumably because bigger areas are needed to satisfy the nutritional requirements of their young. contrary to expectations, Addo males do not travel further and faster on an hourly basis when they are in musth in comparison to when they are not in musth. However, the movement of males in musth is more directional, so that they cover a greater area of the park. Males associate more frequently with females when in musth than when non-musth. Unlike in other populations, non-musth 'male retirement areas' in AENP are not exclusive, as there is spatial and temporal overlap with female ranges, and hence with the ranges of males in musth. It is suggested that the resulting increased frequency of contact between musth and non-musth males may result in elevated mate competition, and this could contribute to the high levels of aggression observed between Addo bulls.

Journal of Indian Veterinary Assocaition Kerala. Journal of Indian Veterinary Association Kerala 7[3], 1-64. 2002.
Ref Type: Journal (Full)

Acharya, B.P., 2002. Intra-specific aggression in male Asian elephants in Similipal Tiger Reserve, Orissa, India. Indian Forester 128, 247-248.

Alex, P.C., 2002. The Musth, the vicious and the rogue elephants - a review. Journal of Indian Veterinary Association Kerala 7, 26-27.

Anderson, J.R., 2002. Gone fishing: tool use in animals. Biologist (London) 49, 15-28.
Abstract: Chimpanzees impress and fascinate us with their use of tools, including twigs to 'fish' for termites and leaves to soak up liquids. But there are many intriguing examples of tool use described across the animal kingdom. Ants use grain to carry honey, and elephants can grip fly switches in their prehensile trunks. Even animals without limbs may use tools.

Chandrasekharan, K., 2002. Sensitive Points of Elephants. Journal of Indian Veterinary Association Kerala 7, 25.

Chandrasekharan, K., 2002. Specific diseases of Asian elephants. Journal of Indian Veterinary Association Kerala 7, 31-34.
Abstract: The earliest writing describing the diseases of elephants in ancient literature said to be the works on "Gajasastra" (Elephantology) written in Sanskrit by authors like Gautama, Narada, Mrigacharma, Rajaputra and Vyasa. "Hasthyayurveda" a legendary book in Sanskrit written by a safe Palakapya deals with some diseases, treatment, desirable and undesirable points of selection, management practices and some mythological aspects on the origin of elephants. The earliest book in English dealing with diseases of elephants seems to be that of W. Gilchrist "A practical treatise on the treatment of diseases of elephants" published in 1848. Later Slym (1873), Sanderson (1878), Steel (1885), Evans (1910), Herpburn (1913), Milroy (1922), Ptaff (1940), Ferrier (1947), Utoke Gale (1974), Chandrasekharan (1979) and Panicker (1985) have documented their findings on the incidence, etiology and control of diseases of Asian elephants.

Cheeran, J.V., Chandrasekharan, K., Radhakrishnan, K., 2002. Tranquilization and translocation of elephants. Journal of Indian Veterinary Association Kerala 7, 42-46.

Cheeran, J.V., 2002. Training and Management of Elephants. Journal of Indian Veterinary Association Kerala 7, 21-23.

Cheeran, J.V., 2002. Elephant facts. Journal of Indian Veterinary Association Kerala 7, 12-14.

Cheeran, J.V., Radhakrishnan, K., Chandrasekharan, K., 2002. Musth. Journal of Indian Veterinary Association Kerala 7, 28-30.

Eggert, L.S., Rasner, C.A., Woodruff, D.S., 2002. The evolution and phylogeography of the African elephant inferred from mitochondrial DNA sequence and nuclear microsatellite markers. Proc R Soc Lond B Biol Sci 269 , 1993-2006.
Abstract: Recent genetic results support the recognition of two African elephant species: Loxodonta africana, the savannah elephant, and Loxodonta cyclotis, the forest elephant. The study, however,#10; did not include the populations of West Africa, where the taxonomic affinities of elephants have been much debated. We examined mitochondrial cytochrome b control region sequences and four microsatellite#10; loci to investigate the genetic differences between the forest and savannah elephants of West and Central Africa. We then combined our data with published control region sequences from across Africa to#10; examine patterns at the continental level. Our analysis reveals several deeply divergent lineages that do not correspond with the currently recognized taxonomy: (i) the forest elephants of Central Africa;#10; (ii) the forest and savannah elephants of West Africa; and (iii) the savannah elephants of eastern, southern and Central Africa. We propose that the complex phylogeographic patterns we detect in African#10; elephants result from repeated continental-scale climatic changes over their five-to-six million year evolutionary history. Until there is consensus on the taxonomy, we suggest that the genetic and ecological#10; distinctness of these lineages should be an important factor in conservation management planning. #10;

Holdo, R.M., Dudley, J.P., McDowell, L.R., (Lungka, G., 2002. Geophagy in the African elephant in relation to availibility of dietary sodium. Journal of Mammalogy 83, 652-664.
Abstract: We studied the use of mineral licks by African elephants (Loxodonta africana) during the dry season in a Kalahari-sand habitat in Hwange National Park, Zimbabwe, to investigate the role of geophagy as a mechanism for supplementing low Na+ levels in browse and natural water supplies. Plant, water, and soil macrominerals were measured to evaluate mineral availability for elephants during the dry season. Elephant behavior was monitored at licks to investigate the intensity of geophagy (measured by number of mouthfuls of soil consumed) in relation to fecal-Na+ loss. Female elephants, which probably had greater requirements than did males because of pregnancy and lactation, consumed more mouthfuls of soil and spent a greater part of their activity budget feeding on soil than did males, suggesting that geophagy may be driven by a nutritional requirement. We found the following consistent with the Na+-supplementation hypothesis: (1) unlike other minerals, Na+ in woody plants and natural water supplies may be inadequate to meet the minimum requirements of elephants during the dry season; (2) soils consumed by elephants differed from other soils primarily in their high Na+ content; (3) intensity of geophagy was negatively correlated with fecal Na+; and (4) elephants in non-Kalahari-sand habitats do not appear to create or use licks, probably because they are able to meet their Na+ requirements from ubiquitous Na+-rich water supplies, which do not occur naturally in Kalahari-sand habitats.

Kumar, G.A., Ghosh, K.N.A., Sreekumaran, T., Chandrasekharan, K., 2002. Reproduction in elephants. Journal of Indian Veterinary Association Kerala 7, 38-40, 48-48.

McDonnell, S.M., Poulin, A., 2002. Equid play ethogram. Applied Animal Behaviour Science 78, 263-290.
Abstract: An ethogram of play behavior among equids was developed. Several key English-language studies on equids were reviewed to derive a preliminary inventory of specific behaviors to be included in the ethogram. Our primary observations were based on a herd of semi-feral Shetland-type ponies kept at New Bolton Center, University of Pennsylvania School of Veterinary Medicine, Kennett Square, PA. Greater than 100 h of direct observation and photo-documentation focused specifically on play in order to identify play behaviors to be added to the preliminary inventory and to obtain detailed descriptions of each behavior. Additionally, these observations were supplemented with photographs obtained during several years of observational study of this herd for other purposes, and with the cumulative equid observational experience and study notes of the principal investigator with other equid species. An initial draft was sent out to 18 equine behavior colleagues for review. A total of 38 individual behaviors classified into four distinct categories were included in the ethogram. These included object play (14 entries), play sexual behavior (3 entries), locomotor play (14 entries) and play fighting (7 entries). All of the behaviors catalogued from direct observation of the herd were also found in the equid literature. The resulting ethogram offers a practical tool as a field guide or reference for quantitative research and other studies of equid play behavior as well as for teaching of equid behavior.

Rajkamal, P.J., Rajeev, T.S., 2002. Training the Mahouts. Journal of Indian Veterinary Association Kerala 7, 24, 30-30.

Rasmussen, L.E.L., Riddle, H.S., Krishnamurthy, V., 2002. Mellifluous matures to malodorous in musth; Mood-altering secretions by excited male elephants smooth out social interactions. Nature 415, 975-976.

Rees, P.A., 2002. Asian elephants (Elephas maximus) dust bathe in response to an increase in environmental temperature. Journal of Thermal Biology 27, 353-358.
Abstract: (1) A captive herd of Asian elephants (Elephas maximus) exhibited dusting  behaviour when the maximum daily temperature exceeded approximately 13°C, and dusting frequency increased directly with the environmental   temperature. (2) Individual animals showed variation in dusting frequency but this was not related to body mass, suggesting that the function of   dusting is not primarily thermoregulatory. (3) Synchronisation in the   timing of dusting behaviour within the herd suggests that it may have a function in the maintenance of social cohesion. (4) The function of  dusting behaviour could not be determined from the data presented, but it  may be involved in skin care, protection from insects or other parasites, temperature control, protection from radiation or some combination of  these.

Schmid, J., 2002. Keeping circus elephants temporarily in paddocks - the effects on their behaviour. Animal Welfare 4, 87-101.

Suedmeyer, W.K. Conditioning programs for transabdominal ultrasound gestational monitoring in an eastern black rhinoceros (Dicerosbicornis michaeli), african elephant, (Loxodonta africana), african lion (Panthera leo), and bornean orangutan (Pongo pygmaeus pygmaeus). Proc Amer Assoc Zoo Vet.  50-52. 2002.
Ref Type: Conference Proceeding

Suprayogi, B., Sugardjito, J., Lilley, R.P.H., 2002. Management of Sumatran elephants in Indonesia: problems and challenges. In: Baker, I., Kashio, M. (Eds.), Giants on Our Hands: Proceedings of the International Workshop on the Domesticated Asian Elephant, Bangkok, Thailand, 5-10 February 2001. FAO Regional Office for Asia and the Pacific (RAPA), Bangkok; Thailand, pp. 183-194.
Abstract: The status of domestic elephants in Indonesia is described. The development of elephant training centres and reasons for their failure to encourage the use of elephants for logging operations are discussed. Problems of stress and other health problems are described. The availability of resources (feed, water and veterinary support) are discussed. Recommendations for improvement are given.

Wielebnowski, N.C., Fletchall, N., 2002. Noninvasive assessment of adrenal activity associated with husbandry and behavioral factors in the North American clouded leopard population. Zoo Biology 21, 77-98.

Barina, A., Reidl, M., Schmid, L., Schutz, R., Bartos, N., Schwammer, H. Development of an Interactive Elephant Feeding System (EFS). A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  246-249. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Ganslober, U. Behavioural Ecology, Social Relationships, Life History and Evolutionary Constraints in Megaherbivores. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  26-31. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Ganswindt, A., Heistermann, M., Hodges, J.K. Faecal Glucocorticoid and Androgen Metabolite Excretion in Male African Elephants (Loxodonta africana). A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  258. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Garai, M. Social Behaviour of the Elephants at Pinnawela Elephant Orphanage, Sri Lanka. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  32-40. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Gsandtner, H., Schwammer, H. Future Perspectives for Elephant-Keeping in Circuses. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  263. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Hart, B., Hart, L., Mccoy, M. Cognition and Tool Use in Elephants. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  41. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Hart, B.L., Hart, L.A., Mccoy, M., Sarath, C.R., 2001. Cognitive behaviour in Asian elephants: use and modification of branches for fly switching. Animal Behaviour 62, 839-847.
Abstract: This article reports on a study which consists of two parts, the first part reports on the observations of one juvenile and 33 adult wild elephants comprising 26.7 h of cumulative observations in Nagarhole National Park, Karnataka, India. Eight of these elephants were observed using branches presumably to repel flies. In the second part of the study, conducted on  the same park, 13 captive elephants, maintained under a naturalistic system, were given branches that were too long or bushy to be effectively used as switches. Eight of these elephants modified the branch on at least one trial to a smaller branch, or modify the branch and switch with the altered branch. The style of branch modification seen in the elephants that did modify was of two general approaches. One was done either of two ways: (1) by holding the main stem on the ground with a front foot and  pulling the side stem off with the trunk; (2) by coiling the trunk around  the side stem and twisting and swishing it against the weight of the main branch.

Hart, L.A., Arnason, B.T., O'Connell-Rodwell, C.E. Bioseismic communication mechanisms in elephants and rhinoceroses. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  42-46. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Heine, N., Kurt, F., Pieler, E., Weihs, W. Social Roles, Family Units and the Formation of Clans in Asian Elephants of the Uda Walawe National Park (Sri Lanka). A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  47-51. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Horwath, A., Kratochvil, H., Schwammer, H. Sounds of a Newborn African Elephant (Loxodonta africana) in Captivity. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  265-267. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Houston, D.C., Gilardi, J.D., Hall, A.J., 2001. Soil consumption by elephants might help to minimize the toxic effects of plant secondary compounds in forest browse. Mammal Review. 31, 249-254.
Abstract: African Elephants Loxodonta africana regularly eat soil. At some sites, such as Mount Elgon on the Kenya-Uganda border, extensive caves have been excavated in the mountainside by their quarrying activity (Redmond, 1984; Bowell, 1992). The material eaten often has little organic content, and this might better be described as a weathered, soft, friable rock, but for convenience we refer to it here as soil. Such soils have been shown to offer sodium, calcium and other mineral nutrients which may be lacking from their normal plant diet (Weir, 1969, 1972, 1973 Moe, 1992; Bowell, 1992; Eksteen & Bornman, 1990). Elephants living in the cloud forest on the eastern escarpment of Ngorongoro in northern Tanzania regularly visit a number of low cliffs to prise away lumps of soil which they eat. This material does not taste "salty" to the human tongue, and we present here an analysis of this material which suggests that oneof its functions may be to assist the animals in the digestion of forest browse through its ability to detoxify the high concentrations of plany secondary compounds found in tropical forest trees.

Kalk, P., Wilgenkamp, C., 2001. Elephant Foot Care Under the Voluntary-Contact System: Problems and Solutions              . In: Csuti, B., Sargent, E.L., Bechert, U.S. (Eds.), The Elephant's Foot. Iowa State University Press, Ames, Iowa, USA, pp. 63-64.
Abstract: Dependable, consistent access to an elephant's nails and pads is a crucial aspect of keeping captive elephants healthy.  This requires the cooperation from the elephant, a facility design that allows safe access to the elephant, and flexible thinking from those who provide the care.  This is true under any method of elephant management, but provides some new challenges with the relatively new, voluntary (or protected) contact style of elephant handling.  We discuss in this chapter some simple elephant facility modifications and training strategies that have allowed us to provide sound foot care for elephants in a voluntary-contact facility. Reliable foot care is critical to the health of captive elephants.  There has been considerable concern about the restricted access to elephant feet, nails and pads under a voluntary-contact system (Priest 1994).  Voluntary contact (VC) is a more accurate and descriptive term than protected contact because it emphasizes the elephants' voluntary cooperation with the keeper (Doherty et al. 1996).  VC has been the sole method of management for four of our Indian Elephants (Elephas maximus indicus) since May 1995.  We describe in this chapter our solutions to obstacles in VC elephant foot access, care, and treatment.  Simple facility modification and systemic training, with thoughtful positioning of the elephant relative to the keeper, has allowed us to provide reliable, sound care for our elephants' feet.

Kam, R., 2001. Preoperative conditioning and postoperative treatments of a protected-contact bull elephant. In: Csuti, B., Sargent, E.L., Bechert, U.S. (Eds.), The Elephant's Foot. Iowa State University Press, Ames, Iowa, USA, pp. 127-131.

Kurt, F., Garai, M. Stereotypies in Captive Asian Elephants - A Symptom of Social Isolation. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  57-63. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

McComb, K., Moss, C., Durant, S.M., Baker, L., Sayialel, S., 2001. Matriarchs as repositories of social knowledge in African elephants. Science 292, 491-494.
Abstract: Despite widespread interest in the evolution of social intelligence, little is known about how wild animals acquire and store information about social companions or whether individuals possessing enhanced social knowledge derive biological fitness benefits. Using playback experiments on African elephants (Loxodonta africana), we demonstrated that the possession of enhanced discriminatory abilities by the oldest individual in a group can influence the social knowledge of the group as a whole. These superior abilities for social discrimination may result in higher per capita reproductive success for female groups led by older individuals. Our findings imply that the removal of older, more experienced individuals, which are often targets for hunters because of their large size, could have serious consequences for endangered populations of advanced social mammals such as elephants and whales.

Pennisi, E., 2001. Behavioral ecology. Elephant matriarchs tell friend from foe. Science 282, 417-419.

Petersen, J.E., Hastings, A., 2001. Dimensional approaches to scaling experimental ecosystems: designing mousetraps to catch elephants. Am. Nat. 157, 324-333.
Abstract: Enclosed experimental ecosystems (mesocosms) are small relative to their natural counterparts, are typically operated for short durations relative to the timescales of a number of important ecological processes, and also often have reduced biological and physical complexity relative to nature. These reductions in time, space, and complexity scales have been cited as sources of unrealistic ecological behavior within mesocosms and raise questions about extrapolating results from mesocosms to nature. Dimensional analysis, a technique widely used by engineers to create scale models, uses compensatory distortion as a means of maintaining dynamic similarity in properties and relationships of interest. Although biological parameters are generally less controllable than physical ones, a variety of dimensional approaches can be taken to maintain such key ecological properties as effective habitat size, environmental variability, vertical and horizontal gradients, interactions among habitats, and control of experimental artifacts. To date, application of dimensional approaches to mesocosm design has been largely intuitive and idiosyncratic. We argue that a more explicit, systematic, and quantitative approach will increase realism and may also provide a critical means of developing, testing, and advancing our understanding of scaling relationships in nature

Pieler, E., Kurt, F. Tool Behavior in Asian Elephants. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  104-109. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Rasmussen, L.E.L., Riddle, H.S. Musth in Teenage Male Asian Elephants (Elephas maximus): The What & the Why of their Chemical Signals. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  110. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Reimers, M. Quantifying Locomotion in African Elephants (Loxodonta africana) at the Vienna Zoo. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  284. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Reimers, M., Schmidt, S., Kurt, F. Daily Activities and Home Ranges in Wild Asian Elephants of the Uda Walawe National Park (Sri Lanka). A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  115-118. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Riddle, H.S., Rasmussen, L.E.L. Are Female African Elephants Messaging through Volatile Chemicals? Studies from European, USA, and African Groups. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  119-120. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Sarma, K.K., 2001. Musth in Asian Elephant. Central Zoo Authority, New Delhi, India.

Schmid, J., Heistermann, M., Ganslosser, U., Hodges, J.K., 2001. Introduction of foreign female Asian elephants (Elephas maximus) into an existing group: behavioural reactions and changes in cortisol levels. Animal-Welfare 10, 357-372.
Abstract: The present study examined the extent to which the introduction of three female Asian elephants (aged 3, 11 and 27 years) into a group of 5 (1 male, 4 female) elephants at Munster zoo, Germany, affects the behaviour and urinary cortisol levels of the animals involved. At Munster, only the females were monitored , while the bull was mainly kept separate. Behavioural observations were carried out before transfer and during the six-month period following transfer. Urine samples were collected regularly from each elephant during the whole observation period. All the elephants showed behavioural changes to the process of introduction. The transferred animals increased their social behaviour after arrival in the foreign zoo. Two of them showed an increase in stereotypies and one a reduction in stereotypes. The elephants at Munster reacted with decreased frequencies of stereotypies and increased frequencies of social behaviour and manipulation/exploration behaviour. Six months after transfer, three of the four elephants at Munster and one of the three transferred elephants showed nearly the same behavioural activity pattern as before transfer. One female still showed elevated stereotypic behaviour. From the four elephants in which cortisol measurements could be reliably performed (two of the transferred elephants and two elephants at Munster), only one individual at Munster responded to the process of introduction with a short-term elevation in urinary cortisol levels. One elephant showed a negative correlation between locomotion and cortisol levels and one a positive correlation between stereotypies and cortisol levels. Taken together, the results suggest that transfer and introduction caused some stress responses in the elephants, but that stress was neither prolonged nor severe. Serious welfare problems may have been prevented through individual behavioural coping mechanisms and former experience with stressful situations.

Schmidt, S., Reimers, M., Kurt, F. Sleep in a Herd of Captive Asian Elephants in the Pinnawela Elephant Orphanage (Sri Lanka). A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  124-128. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Schulte, B.A. Object Manipulation in Captive Asian Elephants. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  288. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Schulte, B.A., Slade, B.E., Rasmussen, L.E.L. The Trunk and Tail of Elephant Communication: Studies on Captive Asian Elephants. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  286. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Schwammer, H., 2001. Elephant Husbandry and Foot Care at the Schonbrunner Tiergarten, Vienna. In: Csuti, B., Sargent, E.L., Bechert, U.S. (Eds.), The Elephant's Foot. Iowa State University Press, Ames, Iowa, USA, pp. 69-71.
Abstract: Zoo visitors consistently rank elephants as number one on their list of the most fascinating animals.  In the wild, the Asian elephant is nearing extinction and the African elephant is endangered.  In captivity, the breeding results are still far from sufficient to maintain the zoo population.  The cause is a lack of zoo facilities able to house bulls.  In addition, many elephant keepers and handlers have insufficient training and knowledge of these species.  There is considerable discussion about the merits of two handling methods: protected contact (hands off) or free contact ( hands on) (Doherty et al. 1996, Priest 1994).  In 1996 the Schonbrunner Tiergarten, Vienna, erected a new facility for African elephants (Pechlaner et al. 1997).  The facility provides for protected-contact or hands-off management for a 7-year old bull and free contact system for the cows.  The latter approach represents the most promising method for managing cows when well-trained animal keepers are available.

Swanepoel, H. African Elephant-Assisted-Therapy for Handicapped Children. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  132. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Wehnelt, S. The New Elephant Exhibit at Chester Zoo - High Husbandry and Welfare Standards. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  293. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Wemelsfelder, F., Hunter, T.E.A., Mendl, M.T., Lawrence, A.B., 2001. Assessing the 'whole animal': A free choice profiling approach. Animal Behaviour 62, 209-220.
Abstract: The qualitative assessment of animal behaviour summarizes the different aspects of an animal's dynamic style of interaction with the environment, using descriptors such as 'confident', 'nervous', 'calm' or 'excitable'. Scientists frequently use such terms in studies of animal personality and temperament, but, wary of anthropomorphism, are reluctant to do so in studies of animal welfare. We hypothesize that qualitative behaviour assessment, in describing behaviour as an expressive process, may have a stronger observational foundation than is currently recognized, and may be of use as an integrative welfare assessment tool. To test this hypothesis, we investigated the inter- and intraobserver reliability of spontanous qualitative assessments of pig, Sus scrofa, behaviour provided by nine naïve observers. We used an experimental methodology called 'free choice profiling' (FCP), which gives observers complete freedom to choose their own descriptive terms. Data were analysed with generalized Procrustes analysis (GPA), a multivariate statistical technique associated with FCP. Observers achieved significant agreement in their assessments of pig behavioural expression in four separate tests, and could accurately repeat attributing expressive scores to individual pigs across these tests. Thus the spontaneous qualitative assessment of pig behaviour showed strong internal validity under our controlled experimental conditions. In conclusion we suggest that qualitative behaviour assessment reflects a 'whole animal' level of organization, which may guide the intepretation of behavioural and physiological measurements in terms of an animal's overall welfare state.

Wilson, M.L., Bloomsmith, M.A., Crane, M., Maple, T.L. Behavior and serum cortisol concentrations of three captive African elephants ( Loxodonta africana): preliminary results. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  147-149. 2001. Vienna, Austria, Schuling Verlag.
Ref Type: Conference Proceeding

Wingate, L., Lasley, B. Is Musth a Reproductive Event: An Examination of Arguments For and Against this View. A Research Update on Elephants and Rhinos; Proceedings of the International Elephant and Rhino Research Symposium, Vienna, June 7-11, 2001.  150-156. 2001. Vienna, Austria, Schuling Verlag. 2001.
Ref Type: Conference Proceeding

Brown, J.L. Zoo Biology. Special Issue on elephant biology 19[5], 1-184. 2000.
Ref Type: Journal (Full)
Abstract: This issue focuses on elephant biology and includes the following topics: ultrasonography of the urogenital tract in elephants Loxodonta africana and Elephas maximus (an important tool for assessing female and male reproductive function); reproductive endocrine monitoring of elephants (an essential tool for assisting captive management); ultrasonography of the oestrous cycle in female African elephants; review of a newly recognized disease of elephants caused by endotheliotropic herpesviruses; tuberculosis in elephants in North America; how chemical signals integrate Asian elephant society; elephant communication; social structure and helping behaviour in captive elephants; a postcryogenic comparison of membrane fatty acids of elephant spermatozoa; and first disclosure and preliminary investigation of a liquid released from the ears of African elephants.

Dale, R.H.I., Jordan, N., Kinnett, S., Beach, L., Noble, J. Behavioral Development of Elephant Calves: Review with Examples from the Indianapolis Zoo. Elephants: Cultural, Behavioral, and Ecological Perspectives; Program and Abstracts of the Workshop.  8. 2000. Davis, CA. 2000.
Ref Type: Conference Proceeding

Fernando, P., Lande, R., 2000. Molecular genetic and behavioral analysis of social organization in the Asian elephant (Elephas maximus). Behavioral Ecology and Sociobiology 48, 84-89.
Abstract: We report on the genetic evaluation and behavioural study of social organization in the Asian elephant (Elephas maximus). Although Asian elephants and African elephants (Loxodonta africana) were previously thought to have similar social organizations, our results demonstrate a substantial difference in the complexity and structure of Asian elephant social groupings from that described for African savanna elephants. Photographic cataloguing of individuals, radio telemetry, and behavioural observations in Ruhuna National Park, Sri Lanka, enabled us to assign associated females and young to four groups with overlapping ranges. Genetic sampling of individuals from the four groups in Ruhuna National Park and three other groups in surrounding areas, conducted through PCR amplification and sequencing of mitochondrial DNA from dung, supported the matriarchal nature of female groups and the lack of inter-group transfer of females. Behaviourally and genetically, the identified social groups were best described as "family groups". We did not find any evidence for the existence of social groups of higher complexity than family groups.

Gruber, T.M., Friend, T.H., Gardner, J.M., Packard, J.M., Beaver, B., Bushong, D., 2000. Variation in stereotypic behavior related to restraint in circus elephants. Zoo Biology 19, 209-221.
Abstract: The effect of penning and chaining on circus elephant behaviour was determined. The behaviours of 10 Asian elephants (Elephas maximus) and 3 African elephants (Loxodonta africana) were videotaped for at least 24 h at each of 6 performance locations. At 4 of these locations, the elephants were restrained in electric-wire pens on turf and at 2 locations, the elephants were restrained via leg chains on macadam. Instantaneous scan sampling at 5-minute intervals during each observation period was conducted to determine the frequency of occurrence of seven behaviour categories: aggression, comfort, ingestion, locomotion, resting, social, and stereotypy. Stereotypies and social interaction (P < 0.0008) were more likely observed with chained restraint. Comfort, ingestion, and locomotion activities were less likely (P < 0.0037) with chained restraint, whereas aggression and resting activities were not affected by the type of restraint (P > 0.6254). Variation in stereotypic activity was related to age (P=0.0001), with younger elephants more likely to show stereotypic activity than older elephants.

Hart, B.L., Hart, L.A., Mccoy, M., Sarath, C.R. Tool Use as a Marker of Cognitive Behavior in Elephants. Elephants: Cultural, Behavioral, and Ecological Perspectives; Program and Abstracts of the Workshop.  12-13. 2000. Davis, CA. 2000.
Ref Type: Conference Proceeding

Kahl, M.P., Armstrong, B.D. Visual displays in wild musth males of the Arican Elephant. Proceedings of the Fifth International Elephant Research Symposium.  2000.
Ref Type: Conference Proceeding

Langbauer, W.R., Jr., 2000. Elephant Communication. Zoo Biology 19, 425-445.
Abstract: Elephants live in a complex society where both long- and short-distance communication play an important role in the ability to locate mates and to maintain intra- and inter-group cohesion. Elephants use a variety of sensory channels in ways both complementary and redundant to achieve this communication, as well as to advertise physiological states, allow reliable assessment of intent, and engage in other behaviors of group living. The majority of long distance communication is probably via infrasonic vocalizations and chemical signals, while vocalizations, chemical signals, and visual and tactile displays all play a role in short distance interactions. While much is known about the general social and behavioral contexts of elephant communication signals, more work needs to be done to elucidate the specific role of many signals. The next critical step in the study of the elephant's vocal repertoire is to collect and categorize the calls of known individuals for later playback experiments to confirm their function. In addition, the way that physiological state affects chemical signals and vice versa is worthy of further study, as is the role of chemical, acoustic, and perhaps seismic communication in long-distance communication. Tactile and visual displays have been qualitatively described, but there is a need to quantifying their role in the dynamic behaviors (such as conflict management) that maintain elephant society. Finally, the way signals from multiple sensory channels interact has been little studied and provides a rich arena for future work.

Lewis, M.H., Gluck, J.P., Petitto, J.M., Hensley, L.L., Ozer, H., 2000. Early social deprivation in nonhuman primates: long-term effects on survival and cell-mediated immunity. Biol Psychiatry 47, 119-126.
Abstract: BACKGROUND: Early differential social experience of non-human primates has resulted in long-term alterations in behavior and neurobiology. Although brief maternal separation has been associated with changes in immune status, the long-term effects on survival and immune function of prolonged early social deprivation are unknown. METHODS: Survival rates were examined in rhesus monkeys, half of which had been socially deprived during their first year of life. Repeated measures of immune status were tested in surviving monkeys (18-24 years old). Peripheral blood T, B, and natural killer lymphocytes, macrophages, and monocytes were measured by flow cytometry. Functional cellular immune activity measures included T-cell proliferative responses to mitogens (concanavalin and phytohemagglutinin), T-cell memory response to tetanus toxoid antigen, T-cell-dependent B-cell proliferative responses to mitogen (PWM) and natural killer cell cytotoxic activity. RESULTS: Despite identical environments following isolation, early social deprivation resulted in a significantly decreased survival rate, males being particularly vulnerable to early death. Early social deprivation was associated with a decrease in the ratio of helper to suppressor T cells, and a significant increase in natural killer cell number and in natural killer cell activity in the surviving monkeys. No differences in T- or B-lymphocyte proliferation following mitogen or tetanus toxoid antigen stimulation were observed. CONCLUSIONS: Prolonged early social deprivation of non-human primates profoundly affected mortality and resulted in lifelong effects on cell-mediated immune status.

McComb, K., Moss, C., Sayialel, S., Baker, L., 2000. Unusually extensive networks of vocal recognition in African elephants. Animal Behaviour 59, 1103-1109.
Abstract: Research on acoustic communication has often focused on signaling between territorial individuals or static neighboring groups. Under these circumstances, receivers have the opportunity to learn to recognize the signals only of the limited number of conspecifics with which they are in auditory contact. In some mammals, however, social units move freely with respect to one another and range widely, providing individuals with opportunities to learn to recognize the signals of a wide range of conspecifics in addition to those of their immediate neighbors. We conducted playback experiments on African elephants, Loxodonta africana, in Amboseli National Park, Kenya, to determine the extent to which adult female elephants, which have a highly fluid social system, can recognize others in the population through infrasonic contact calls. Female elephants could distinguish the calls of female family and bond group members from those of females outside of these categories; moreover, they could also discriminate between the calls of family units further removed than bond group members, on the basis of how frequently they encountered them. We estimated that subjects would have to be familiar with the contact calls of a mean of 14 families in the population (containing around 100 adult females in total), in order to perform these discriminations. Female elephants thus appear to have unusually extensive networks of vocal recognition, which may prove to be typical of long-lived species that have both fluid social systems and the means for long-distance vocal communication.

Mikota, S.K. Sumatra's elephant training centers: a call for assistance. AAZV and IAAAM Joint Conference.  127-129. 2000.
Ref Type: Conference Proceeding

Mooring, M.S., Benjamin, J.E., Harte, C.R., Herzog, N.B., 2000. Testing the interspecific body size principle in ungulates: the smaller they come, the harder they groom. Animal Behaviour 60, 35-45.
Abstract: Tick removal grooming may be centrally regulated by an internal timing mechanism operating to remove ticks before they attach and engorge (programmed grooming model) and/or evoked by cutaneous stimulation from tick bites (stimulus-driven model). The programmed grooming model predicts that organismic and environmental factors that impact the cost-benefit ratio of grooming (e.g. body size and habitat) will influence the rate of tick removal grooming. The body size principle predicts that smaller-sized animals, because of their greater surface-to-mass ratio, should engage in more frequent tick removal grooming than larger-bodied animals in order to compensate for higher costs of tick infestation. The body size principle may be tested intraspecifically between young and adult animals, or interspecifically among species of contrasting body sizes. To rigorously test the interspecific body size prediction, we observed the programmed grooming (oral and scratch grooming) of 25 species (or subspecies) of bovids at a tick-free zoological park in which stimulus-driven grooming was ruled out. Multiple correlation analysis revealed highly significant negative correlations between species-typical mass and mean species grooming rates when habitat was controlled for in the model. Species-typical habitat type (classified along a gradient from most open to most closed) was positively correlated with mean oral grooming rate, indicating that species tended to groom at a higher rate in woodland and forest habitats (where typical tick density would be high) compared with more open environments. Species mass accounted for up to two-thirds of the variation in grooming rate across species, whereas habitat accounted for ca. 20% of variation in oral grooming. Similar results were obtained when the analysis was expanded to include 36 species/subspecies of six different families. The body size principle can therefore account for a large proportion of species-typical differences in programmed grooming rate among ungulates. However, to understand the tick defense adaptations of very large mammals that rarely or never engage in oral or scratch grooming (e.g. elephants, giraffes, rhinoceros), alternative tick defense strategies must be considered, such as thick skin, wallowing, rubbing and tolerance of oxpeckers and other tick-eating birds.

Rasmussen, L.E.L. Wild and Non-wild Elephants: How Two Modes of Olfaction and a Multitude of Chemical Signals and Pheromones Influence Elephant Behavior. Elephants: Cultural, Behavioral, and Ecological Perspectives; Program and Abstracts of the Workshop.  19-20. 2000. Davis, CA. 2000.
Ref Type: Conference Proceeding

Rasmussen, L.E.L., Krishnamurthy, V., 2000. How chemical signals integrate Asian elephant society: the known and the unknown. Zoo Biology 19, 405-423.
Abstract: The importance of chemical senses to elephants was recognized in anecdotal observations by ancient humans. Modern scientific tools, such as molecular biological techniques, highly sensitive gas chromatographic/mass spectrometric instrumentation, and statistically valid ethological methods, have allowed the study of real events of chemical communication between elephants. Such communication encompasses long- and short-range navigation, relationship recognition, and inter- and intra-sexual exchange of reproductive condition, metabolic state, and social status. Asian elephants emit large amounts of complex chemical mixtures in breath and urine, and in secretions from the temporal gland, inter-digital glands, and ears. Some emitted chemicals originate in blood and may be metabolic products; others are secretory products, at times apparently under hormonal control. The wide variety of emitted compounds includes hormones, proteins, and volatile compounds; selected volatile ketones and an acetate ((Z)-7-dodecenyl acetate) function as chemical signals and a pheromone, respectively. Some of these specific chemicals identified in emissions from Asian elephants dwelling in the United States have been found to be present in the exudates from elephants in India. This similarity is demonstrable for three metabolic conditions: pregnancy in females and pre- and post-musth males. Future chemical communication studies on male elephants should focus on musth and its relevance to reproduction and male social structures. Such investigations should include hormones, metabolites, brain chemistry, and possible primer pheromones. For females, the factors influencing possible estrous synchrony, what role primer pheromones play in female reproduction, how chemical signals influence social behavior, and whether luteinizing hormone influences pheromone production are among remaining fundamental questions.

Riddle, H.S., Riddle, S.W., Rasmussen, L.E.L., Goodwin, T.E., 2000. First disclosure and preliminary investigation of a liquid released from the ears of African elephants. Zoo Biology 19, 475-480.
Abstract: This report is the first documentation, both behaviorally and chemically, of a phenomenon observed among African elephants (Loxodonta africana) whereby a sudden, often stream-like discharge of liquid is seen from the auricular orifice.  During this initial investigation, multiple samples of the fluid have been collected for analysis of physical properties and components.  Trace organic chemicals which are apparently of elephant origin have been identified in the ear liquid, and the aqueous nature of the liquid has been demonstrated.  The continuing objectives of this work and related studies are to determine the specific source of the liquid with particular focus on a search for auricular glands, to further characterize potential conspecific chemical signals, and to document more precisely particular social situations when this phenomenon occurs.

Schulte, B.A., Feldman, E., Lambert, R., Oliver, R., Oliver, R., 2000. Temporary ovarian inactivity in elephants: relationship to status and time outside. Physiol Behav 71, 123-131.
Abstract: The captive elephant population in North America is in reproductive decline and,without importation from the wild, may cease to be viable within the next several decades. The estrous cycle of three captive, reproductive-age African elephants was monitored for 3 years by measuring serum progesterone concentrations. Each elephant experienced one or more episodes of extended low progesterone (>12 weeks), analogous to supposed terminal cessation of estrous cyclicity or 'flatlining' that has been described in some captive Asian and
African elephants. Other studies have reported lengthy non-luteal (follicular)phases that indicate extended episodes of ovarian inactivity; however, this phenomenon has not been examined in detail. In this study, total duration of temporary ovarian inactivity or acyclicity followed a social rank pattern, with the most subordinate female having the longest and the dominant female the shortest duration. During periods of acyclicity, the number of hours the elephants spent outside was significantly less than during non-luteal or luteal phases of the cycle. Except in one instance, behavioral data recorded by elephant keepers during their interactions with the elephants showed no change in handling during periods of ovarian inactivity. Further study is necessary to distinguish the causative agent for temporary cessation of estrous cyclicity. Understanding this phenomenon is imperative for the future reproductive viability of captive elephant populations.

Schulte, B.A., 2000. Social structure and helping behavior in captive elephants. Zoo Biology 19, 447-449.
Abstract: Selective pressures on individual behavior serve to mold social structure and subsequent levels of cooperation in social species, including elephants.  Adult Asian (Elephas maximus) and African (Loxodonta africana) elephants have complex social societies that differ markedly between the sexes.  Elephant social structure and cooperative behavior may have evolved in part to prepare young elephants to meet the challenges of an expansive environment, predators and parasites.  In captivity, such forces are relaxed or eliminated, yet captive elephants display a similar repertoire of behaviors as their wild counterparts.  Elephants live in herds of related adult females from several generations. This female social unit is centered about calves and the matriarch.  Adult males reside in bachelor herds, singly, or frequent female groups, depending upon male age and reproductive condition.  In captivity female elephants are usually housed together although group size is smaller than in the wild, while adult males are generally housed separately from other elephants.  Beyond these general similarities between wild and captive settings, captive elephants are housed in a wider range of group sizes, degrees of relatedness and age structures than would be typical in the wild.  This variation in social structure can be used to explore questions about cooperative behavior.  Following a comparison of wild and captive social structures for elephants, three such potential avenues for study are discussed.  Specifically, research with captive elephants could shed light on the development of social behavior, the functional significance of allomothering and the ability of elephants to recognize kin or social group members.  Beside the potential theoretical implications and insight into wild elephant behavior, the examination of social behavior also has important ramifications for the management of animals in captivity.

Simonet, P., Krishnamurthy, R. Self-recognition Among Captive Asian Elephants: Preliminary Results and Future Studies. Elephants: Cultural, Behavioral, and Ecological Perspectives; Program and Abstracts of the Workshop.  22. 2000. Davis, CA. 2000.
Ref Type: Conference Proceeding

Slotow, R., van Dyk, G., Poole, J., Page, B., Klocke, A., 2000. Older bull elephants control young males. Nature 408, 425-426.

Stoinski, T.S., Daniel, E., Maple, T.L., 2000. A preliminary study of the behavioral effects of feeding enrichment on African elephants. Zoo Biology 19, 485-493.
Abstract: Although commonly practiced in many species, feeding enrichment in elephants is understudied. The behavioural effects of feeding enrichment were tested in 3 African elephants by substituting an equal dry weight of browse for hay in an ABA design. A significant increase in feeding was observed when the browse was present. In contrast, drinking and inactivity decreased. Additionally, changes in feeding, inactivity, and time spent in contact were observed outside the time the browse was actually present. The relationship of these changes to the experimental methodology was unknown. A significant increase in visibility to zoo visitors during the browse conditions demonstrated that browse is an effective naturalistic method for increasing visibility as well as species-typical behaviours. Managers of captive elephants should consult with a nutritionist to address issues of energetics, nutritional content, and secondary compounds when using browse as a feeding enrichment.

Wingate, L., Lasley, B. The Significance of Musth in Bull Elephants: Is It a Reproductive Event? Elephants: Cultural, Behavioral, and Ecological Perspectives; Program and Abstracts of the Workshop.  24. 2000. Davis, CA. 2000.
Ref Type: Conference Proceeding

Brockett, R.C., Stoinski, T.S., Black, J., Markowitz, T., Maple, T.L., 1999. Nocturnal behavior in a group of unchained female African elephants. Zoo Biology 18, 101-109.
Abstract: A study of 3 unchained female African elephants was undertaken to document their nocturnal behaviour. The subjects were observed between the hours of 1800 and 0800 for 10 weeks in the summer of 1992 (total of 172 h) and 14 weeks in the summer of 1994 (total of 153 h). Scan data were collected every 5 min to gather information on activity budgets, social proximity and space utilization. All-occurrence data were collected on social and non-social behaviours. In each year of the study, the subjects spent equivalent amounts of time eating, lying, standing, and walking. Additionally, subjects spent half of their time within 1 body length of another animal and utilized all 3 available enclosures. Social and non-social behaviours were frequent, and these data plus the activity profiles revealed the elephants generally were most active between the hours of 1800 and 2400 and 0600 and 0700. The findings suggest that the use of no restraints is currently an effective strategy for this elephant group. The high activity levels observed during many of the early evening hours suggest that zoos could permit increased activity and social interactions by extending the hours when the elephants are unchained.

Durrheim, D.N., Durrheim, D.N., 1999. Risk to tourists posed by wild mammals in South Africa. J Travel Med 6, 172-179.
Abstract: BACKGROUND: One of South Africa's principal tourist attractions is the opportunity to encounter Africa's large mammals in the wild. Attacks by these mammals can be exceptionally newsworthy with potentially deleterious effects on tourism. Little is known about the risk of injury and death caused by wild mammals to visitors to South Africa's nature reserves. The aim of this study was to determine the incidence of fatal and nonfatal attacks on tourists by wild mammals in South Africa and to ascertain avoidable factors, if any. METHODS: Commercial press records covering all South African Newspapers archived at the Independent Newspapers' central library were systematically reviewed for a 10-year period, January 1988 to December 1997 inclusive, to identify all deaths and injuries to domestic and international tourists resulting from encounters with wild mammals in South Africa. All of these incidents were analyzed to ascertain avoidable factors. RESULTS: During the review period seven tourists, including two students from Thailand and a German traveler, were killed by wild mammals in South Africa. Three of the four deaths ascribed to lions resulted from tourists carelessly approaching prides on foot in lion reserves. A judicial inquiry found that the management of a KwaZulu-Natal Reserve was culpable for the remaining death. Tourist ignorance of animal behavior and flagrant disregard of rules contributed to the two fatalities involving hippopotami. The unusual behavior manifested by the bull elephant responsible for the final death, resulted from discomfort caused by a dental problem to this pachyderm. During the same period there were 14 nonfatal attacks on tourists, including five by hippo, three by buffalo, two by rhino, and one each by a lion, leopard, zebra and musth elephant. Only the latter occurred while the visitor was in a motor vehicle. Tourist ethological naivete and failure to determine the experience of trail guides prior to travel, resulted in inadvertent agonistic behavior, unnecessary risk-taking and avoidable injury. CONCLUSIONS: This retrospective study has shown that attacks on tourists by wild mammals in South Africa are an uncommon cause of injury and death. Sensible precautions to minimize this risk include remaining in a secure motor vehicle or adequately fenced precincts while in the vicinity of large mammals, rigidly observing nature reserve instructions, never approaching animals that appear ill, malnourished, displaying aggressive behavior traits or female wild mammals with young, and demanding adequately trained and experienced game rangers when embarking on walking trails. Any behavior that might be construed as antagonistic and which could provoke an attack by large mammals should be avoided (e.g., driving directly at a lion). Visitors need to be informed of classic signs of aggression, in particular in elephants, which will allow timely avoidance measures to be taken. The risk-enhancing effect of excessive alcohol intake is undesirable in the game reserve setting, as is driving at high speed after dusk in areas where hippos graze. Local advice on personal safety in wildlife reserves and the credentials of trail guides should be obtained from lodge or reserve management, tourism authorities or the travel industry prior to travel to game reserves.

Friend, T.H., Parke, M.L., 1999. The effect of penning versus picketing on stereotypic behavior of circus elephants. Applied Animal Behaviour Science 64, 213-225.
Abstract: The behaviour of 9 female Asian elephants (Elephas maximus) who performed 2 shows each day with a circus that travelled to a new location in the USA (40 to 250 km from the previous day's lot) daily or every 2 days was studied during the 1996 and 1998 seasons. When not performing or working, the elephants were picketed during 1996 in the traditional fashion. During the 1998 season, the same elephants were maintained exclusively in small (approximately 52 m2 per elephant) portable electric pens. Camera and time-lapse recorders were used to videotape the behaviour of each elephant, while picketed or penned, for three _24 h periods during the 1996 and 1998 seasons. The behaviour of each focal elephant was recorded at 5 min intervals. The amount of time the elephants spent stereotypic weaving was significantly decreased by keeping the elephants in pens when compared to picketing. The incidence of all stereotypic behaviour (weaving, head bobbing and trunk tossing) was also significantly decreased when the elephants were kept in pens. The total amount of time spent performing all stereotypic behaviours (weaving, head bobbing and trunk tossing) was negatively correlated with age when the elephants were picketed in 1996 and somewhat less correlated with age when penned in 1998. Time spent performing all stereotypic behaviour was not correlated with time spent eating or time spent lying when the elephants were picketed or penned. It is concluded that portable electric pens are preferred over picketing because the elephants show reduced stereotypic behaviour, they appear to be calmer when out of the pens for work or performances, and they can be kept cleaner.

Friend, T.H., 1999. Behavior of picketed circus elephants. Applied Animal Behaviour Science 62, 73-88.
Abstract: The behaviour of 14 female Asian (Elephas maximus) and 3 female African (Loxodonta africana) elephants who performed 2 shows each day with a circus that travelled to a new location (40 to 250 km from the previous day's lot) daily, was studied. When not performing or working, the elephants were picketed (chained) in 2 separate groups of 8 and 9 elephants each. While they were picketed, a separate camera and time-lapse recorder videotaped the behaviour of members of each group for 4, 8 h periods during the 1995 season, and 3, 24 h periods during 1996 season. The behaviour of each member of the focal groups was recorded at 5 min intervals. In 1995, stereotypic weaving accounted for 15.9% and head bobbing accounted for 2.8% of observations in 8 of the elephants. In 1996, stereotypic weaving accounted for 14.3% and head bobbing accounted for 2.9% of observations in 12 of the elephants. The behaviour of individual picketed elephants in 1995 was highly correlated with their behaviour patterns in 1996, despite season and location differences. The elephants spent an average of 33±1.2% of observations eating in 1996. Stereotypic behaviour increased in the 15 min period immediately before water, performances and hay, when compared with their frequency during the 3 preceding 15 min periods indicating 'anticipation' of water and performances, and a lack of substrate to manipulate or eat in regards to hay. Time the elephants spent off the picket line (performing, working, giving rides, etc.) tended to be negatively correlated with weaving.

Karunaratne, S.H.P.P., Ranawana, K.B., 1999. A preliminary study on feeding activity patterns and budgets of domesticated elephants (Elephas maximus maximus) in Sri Lanka. Ceylon Journal of Science, Biological Sciences 27, 61-65.
Abstract: Feeding activity patterns of 3 domesticated elephants were studied for 7 consecutive days. Four food items (I) jak (Artocarpus heterophyllus) leaves and twigs, (II) a browse mixture (excluding jak), (III) Panicum maximum and (IV) a grass mixture (excluding Panicum maximum) were offered separately and feeding activities were observed for 5 each day. Although P. maximum and grass mixture were consumed in large quantities, all 3 elephants spent most of their feeding time on jak and browse mixture (65.4-79.8% of their feeding time). This was mainly because they had a special preference for feeding on the bark of the large twigs of jak and browse and the peeling process was very time-consuming. It is apparent that although grass acts as a filling food, the bark of jak and browse branches serve some specific need of the elephant.

Mohan, A.B., Lakshmi, B.B., 1999. The successful capture and training of two strayed wild elephants "Jay - Vijay" in Sri Venkateswara Wildlife Sanctuary, Andhra Pradesh. Zoos' Print Journal 14, 1-6.

Morimura, N., Ueno, Y., 1999. Influences on the feeding behavior of three mammals in the Maruyama Zoo: bears, elephants, and chimpanzees
532. J. Appl. Anim Welf. Sci. 2, 169-186.
Abstract: Modifications of feeding conditions are essential to the establishment of environmental enrichment in zoos. In this study, we attempted to increase the duration of feeding by varying feeding conditions, such as the spatial or temporal distribution of foods, while keeping the sorts and amounts of food the same. Subjects included nonhuman animals reared at the Maruyama Zoo in Sapporo, Japan: 3 bears (Ursus arctos), 2 elephants (Elephas maximus), and 5 chimpanzees (Pan troglodytes). Considering that the feeding ecologies of these animals differed a great deal from those in the wild, we used the focal animal sampling method. Consisting of the spatial dispersive and massed feeding conditions, the experiment with bears found feeding time increased more in the dispersive condition. In addition, the behavioral rhythm of alternate feeding and sleeping appeared. The experiment with elephants, consisting of the temporal dispersive and massed condition, increased feeding time under the massed condition. The experiment using chimpanzees reversed conditions used in the elephant study and increased feeding time under the dispersive condition. Results indicated that these simple modifications both influenced an increasing duration of feeding and affected behavioral patterns in a day

Poole, J.H., 1999. Signals and assessment in African elephants: evidence from playback experiments. Animal Behaviour 58, 185-193.
Abstract: A series of playback experiments using two elephant vocalizations, the 'musth rumble' and the 'oestrous call', was carried out in Amboseli National Park to examine signaling and assessment in African elephants, Loxodonta africana. In response to the musth rumble of a high-ranking male other musth males approached the speaker aggressively, whereas nonmusth males walked away from the stimulus. The call of an oestrous female, too, attracted musth males who approached the speaker rapidly, while nonmusth males listened and then walked away. Females listened and often showed considerable interest in the musth rumbles of males, approaching the speaker and sometimes responding by vocalizing and or secreting from the temporal glands. The experiments bear out earlier observational data and game theory predictions which suggest that by being in or out of musth a male may be conveying information about the relative value he places on contesting his dominance rank and his access to oestrous females. When not visibly in musth, a male may be indicating his intention not to contest access to oestrous females.

Price, E.O., 1999. Behavioral development in animals undergoing domestication. Applied Animal Behaviour Science 65, 245-271.
Abstract: The process of domestication involves adaptation, usually to a captive environment. Domestication is attained by some combination of genetic changes occurring over generations and developmental mechanisms e.g., physical maturation, learning.triggered by recurring environmental events or management practices in captivity that influence specific biological traits. The transition from free-living to captive status is often accompanied by changes in availability and or accessibility of shelter, space, food and water, and by changes in predation and the social environment. These changes set the stage for the development of the domestic phenotype. Behavioral development in animals undergoing domestication is characterized by changes in the quantitative rather than qualitative nature of responses. The hypothesized loss of certain behavior patterns under domestication can usually be explained by the heightening of response thresholds. Increases in response frequency accompanying domestication can often be explained by atypical rates of exposure to certain forms of perceptual and locomotor stimulation. Genetic changes influencing the development of the domestic phenotype result from inbreeding,  genetic drift, artificial selection, natural selection in captivity, and relaxed selection. Experiential contributions to the domestic phenotype include the presence or absence of key stimuli, changes in intraspecific aggressive interactions and interactions with humans. Man's role as a buffer between the animal and its environment is also believed to have an important effect on the development of the domestic phenotype. The domestication process has frequently reduced the sensitivity of animals to changes in their environment, perhaps the single-most important change accompanying domestication. It has also resulted in modified rates of behavioral and physical development. Interest in breeding animals in captivity for release in nature has flourished in recent decades. The capacity of domestic animals to survive and reproduce in nature may depend on the extent to which the gene pool of the population has been altered during the domestication process and flexibility in behavioral development. ''Natural'' gene pools should be protected when breeding wild animals in captivity for the purpose of reestablishing free-living natural populations. In some cases, captive-reared animals must be conditioned to live in nature prior to their release.

Rasmussen, L.E.L., 1999. Evolution of chemical signals in the Asian elephant, Elephas maximus: behavioural and ecological influences. Journal of Biosciences 24, 241-251.
Abstract: In antiquity, the Asian elephant, Elephas maximus, gradually spread southward and eastward to become a successfully surviving, ecologically dominant megaherbivore in the tropical environment of south-east Asia. The changing physical environment forced dynamic fluxes in its social structure and altered its metabolism. Such events shaped the production and ultimately the stability of certain chemicals released by body effluvia. Some of these chemicals took on significance as chemical signals and/or pheromones. This article demonstrates by experimental and observational evidence, and hypothesizes based on speculative reasoning, how and why specific chemical signals evolved in the modern Asian elephant. Evidence, including the functional criteria required by elephant social structure and ecology, is presented for the hypothesis that the recently identified female-emitted, male-received sex pheromone, (Z)-7-dodecenyl acetate evolved first as a chemical signal. Subsequently, the cohesiveness and harmony of small, matriarchally-led female groups were strengthened by a female-to-female chemical signal, recently defined behaviourally. The looser societal structure of freer, roaming males also became bounded by chemical signals; for the males, breath and temporal gland emissions, as well as urinary ones function in chemical signaling. Basic knowledge about elephant chemical signals is now linking chemical information to behaviour and beginning to demonstrate how these signals affect elephant social structure and enable the species to cope with environmental changes.

Sarma, K.K., 1999. Bizarre behaviour of an elephant during xylazine anaesthesia. Indian Veterinary Journal 76, 1018-1019.

Schulte, B.A., Rasmussen, L.E., 1999. Signal-receiver interplay in the communication of male condition by Asian elephants. Animal Behaviour 57, 1265-1274.
Abstract: Signal design and meaning are dependent on the condition of the sender and receiver as well as the response of the receiver. This study examined (1) whether female Asian elephants, Elephas maximus, can distinguish between a conspecific male in musth and nonmusth states using urinary signals, (2) how the oestrous condition of the female affects discrimination, and (3) correlation of female responses with the testosterone level of the male. Musth is a rut-like state displayed by healthy adult male elephants. Males in musth dominate nonmusth males and may be preferred by females as mates. Urine was collected from two captive male Asian elephants during nonmusth periods and from one of these males during times of musth. Samples of musth and nonmusth urine and control liquids were placed in an elephant enclosure weekly for 16 weeks, the length of a female oestrous cycle. Primary response behaviours were approach and four trunk-tip motions, namely sniff, check, place and flehmen. Musth urine consistently elicited greater responses than nonmusth and control samples. Females were more responsive during their follicular (sexually  receptive) than luteal (unreceptive) stages of oestrus. Furthermore, females appeared to be sensitive to the degree of musth as responses increased with rising serum testosterone levels of the male donor. Chemical signals from males are a likely source of honest signals related to status and reproductive condition. Female elephants appear capable of detecting differences in a male based upon urinary chemosignals.

Schulte, B.A., Rasmussen, L.E.L., 1999. Musth, sexual selection, testosterone and  metabolites. In: Johnston, R.E., Muller-Schwarze, D., Sorenson, P.W. (Eds.), Advances in Chemical Communication in Vertebrates 8. Kluwer Academic/Plenum Press, pp. 383-397.
Abstract: Musth is an annual, yet asynchronous, rut-like condition that is experienced by many adult African and Asian male elephants. Behaviorally, musth is characterized by heightened aggression, decreased feeding, urine dribbling, temporal gland secretion and enhanced sexual activity. Musth improves the access of a male to reproductively active females through increased mobility and a higher dominance ranking (intrasexual competition). Whether females prefer males in musth as mates is as yet uncertain (intersexual choice). Females can distinguish among the odors of males in musth and nonmusth. Although behavioral musth has been associated with greatly elevated plasma testosterone levels, a recent study in Sri Lanka shows that intensified aggressiveness follows maximal testosterone secretion and proposes that behavioral musth is a consequence of declining androgen levels. Our data from an Asian male elephant in North America suggest that either declining or rising serum testosterone may be related to "musth behaviors." Our report demonstrates that certain aspects of body physiology are greatly altered during musth. Rather than a single state, our data suggest that musth is an ever-changing condition with some typical stages. Specific chemical compounds released at different stages of musth may serve individually or in combination as honest signals of male condition.

Shepherdson, D., 1999. Environmental enrichment for elephants:current status and future directions. Journal of the Elephant Managers' Association 10, 69-77.

Managing giants.  1998. Chicago, Ill. USA, Chicago]: Lincoln Park Zoo,  1 videocassette (17 min.).
Ref Type: Online Source
Abstract: Keeper and animal safety are emphasized in the Lincoln Park Zoo's "Elephant  Management Program." Training of both the animals and keepers are described

Karunaratne, S.H.P.P., Ranawana, K.B., 1998. A preliminary study of the food preference of domesticated elephants (Elephas maximus L) in Sri Lanka. Ceylon Journal of Science, Biological Sciences 25, 58-64.
Abstract: Food preference of 3 domesticated elephants were studied for 7 days. Four food items, jack (Artocarpus heterophyllus) leaves and twigs, a browse mixture (excluding jack), Panicum maximum and a grass mixture (excluding P. maximum) were offered in separate bundles. Elephants were allowed to feed on these for 24 h and at the end of each feeding trial, food items were separated into 4 groups and weighed. On the basis of the amount consumed and discarded, food preference was calculated using Manly's Alpha index. All 3 elephants used in the experiment showed a preference for P. maximum or the grass mixture.

Payne, K., 1998. Silent Thunder : in the presence of elephants. Simon & Schuster, New York.

Rasmussen, L.E.L., Schulte, B.A., 1998. Chemical signals in the reproduction of Asian (Elephas maximus) and African (Loxodonta africana) elephants. Animal Reproduction Science 53, 19-34.
Abstract: Asian (Elephas maximus) and African (Loxodonta africana) elephants exhibit polygynous mating that involves female choice of mates and male-male competition for access to females. Chemical signals mediate intersexual and intrasexual interactions associated with reproduction. The need for reliable and honest signals is accentuated by the markedly different social structure of adult males and females. Adult female elephants live in matriarchal herds consisting of a dominant female and several generations of offspring. Adult males are solitary or travel with other males except during breeding periods. Because females have a long 16-week oestrous cycle with a brief 1-week receptive period and a 4-5 year interval between births, a sexually active female is a limited resource. Asian elephant females advertise a forthcoming ovulation by releasing (Z)-7-dodecen-1-yl acetate in their urine during the preovulatory period. African elephants probably produce a sex pheromone as well. Females regularly contact the ano-genital region of other females and show heightened chemosensory responsiveness to urine during the follicular phase. The physiological impacts of this ability to detect reproductive condition (e.g. possible synchronizing or suppressing of oestrus) are uncertain. Males experience an annual period of heightened aggressiveness and highly elevated testosterone concentrations known as musth. Males secrete fluid copiously from their temporal gland and dribble strongly odoriferous urine during musth. Females appear to prefer musth males as mates, and captive Asian females exhibit greater chemosensory responses to urine from males in musth than not. Males in musth are competitively dominant to all other males, even those larger than themselves. Nonmusth males avoid males in musth, and captive Asian bulls show greater interest in musth than nonmusth urine. In captivity subordinate Asian females back away from musth secretions, and females with calves sometimes display protective behaviour. Clearly, chemical signals play an important role in communication by elephants between and within the sexes. Further work is needed to identify more of these chemical messengers and to understand their complete function in mediating reproductive interactions in the elephant social system.

Sai, D.J., Chen, Z.B., Zhang, J.Z., 1998. Study on the courtship-mating behavior of Asian elephant. Chinese Journal of Zoology 33, 28-31.

Schulte, B., 1998. The benefits of behavioral research to elephants and people. Journal of the Elephant Managers Association 9, 216-221.

Stuwe, M., Abdul, J.B., Nor, B.M., Wemmer, C.M., 1998. Tracking the movements of translocated elephants in Malaysia using satellite telemetry. Oryx 32, 68-74.
Abstract: Malaysia incurs damage worth millions of dollars annually from elephants feeding in plantations. In response, the Malaysian Wildlife Department has translocated over 392 elephants from plantations into protected areas since 1974. Dense rain forest and steep terrain at the release sites have made it near impossible to follow the movements of the released elephants and evaluate the success of the programme. In October 1995, a translocated female elephant was fitted with a satellite transmitter to determine its post-release movements. By August 1996, the female had moved within a range of 7000 sq km. The movements were erratic initially and covered a large area. They were followed by a shift in location twice, each time covering successively smaller areas. Attempts to locate and follow the female on the ground revealed no evidence that she travelled in a group. A second, male, elephant was equipped with a satellite transmitter in February 1996 and by August 1996, it had moved within a range of 350 sq km. An attempt to locate and follow the male on the ground provided evidence that he was in a group of at least three individuals. The difference in the behaviour of the two elephants, released at exactly the same location, is striking, but additional elephants need to be followed to increase the sample size and allow a scientific explanation of the findings.

Taylor, V.J., Poole, T.B., 1998. Captive breeding and infant mortality in Asian elephants:  a comparison between twenty Western zoos and three Eastern elephant centers. Zoo Biology 17, 311-332.
Abstract: A questionnaire was designed to assess the importance of reproductive behaviour and husbandry factors on breeding success in captive Asian elephants (Elephas maximus). This was circulated to zoos in Europe and North America in 1996. Data from 20 zoos were analysed. Data were also obtained from 3 elephant centres in Asia (Pinnawela Elephant Orphanage in Sri Lanka, Myanma Timber Enterprise in Myanmar and the Tamilnadu Forest Department in India). The aims were to compare Asian elephant breeding success, establish possible causes for any differences, and make recommendations for improving the welfare and breeding success of the animals. Breeding success in most of the zoos was notably lower and the percentages of stillbirths and infant mortality were relatively higher when compared with those of the centres in Asia. Female elephants in zoos appeared to reach sexual maturity and reproduce earlier than those in the Asian establishments. However, zoo elephants produced fewer young per female. The different facilities and husbandry methods used are described. Recommendations are made for both short- and long-term changes that could be used to modify existing practices to improve the welfare and breeding success of captive Asian elephants.

Wiedenmayer, C., 1998. Food hiding and enrichment in captive Asian elephants. Applied Animal Behaviour Science 56, 77-82.
Abstract: Asian elephants (Elephas maximus) kept in zoos usually spend less time foraging than their wild conspecifics. In order to increase foraging in a group of 5 captive Asian elephants, peanuts were hidden above a distinctive outdoor enclosure structure, the stone border. It was expected that the elephants would learn this association and increase food searching in the outdoor enclosure. After training, the elephants searched more above the stone borders than during baseline, which indicates that they learned the relationship between stone borders and peanuts. But hiding food did not enhance searching behaviour. Therefore, this method of food hiding did not represent an environmental enrichment.

Dehnhardt, G., Friese, C., Sachser, N., 1997. Sensitivity of the trunk of Asian elephants for texture differences of actively touched objects. Zeitschrift fuer Saeugetierkunde 62, 37-39.

Dickerman, R.D., Zachariah, N.Y., Fouraker, M., McConathy, W.J., 1997. Neuroendocrine-associated behavioral patterns in the male Asian elephant (Elephas maximus). Physiology and Behavior 61, 771-773.
Abstract: Steroid-responsive behaviors have been reported in various species; however, the reports thus far on the male Asian elephant (bull) during musth are few in number and most have been conducted on single captive animals for short time periods. The purpose of this investigation was to perform a longitudinal study on steroid-responsive behavior in 3 male Asian elephants from a captive herd of 11 male Asian elephants in Nepal. Male Asian elephants were 18, 25, and 43 years old. The animals had serum collected for 11 months and were observed on a daily basis for aggressive behavior according to the Species Survival Plan (SSP) collection protocol on SSP data sheets. Testosterone (T) and dihydrotestosterone (DHT) were measured in each animal by radioimmunoassay. Testosterone levels rose during musth 26-fold compared to nonmusth, and DHT was elevated 12-fold in musth. Maximal aggressive behavior episodes occurred during peak elevations of T and DHT, with correlation coefficients of 0.82 and 0.89, respectively. Therefore, we suggest that the aggressive episodes are dependent on elevated circulating androgens acting on androgen-responsive neural tissues.

O'Connell, C.E., Arnason, B.T., Hart, L.A., 1997. Seismic transmission of elephant vocalizations and movement. Journal of the Acoustical Society of America 102, 3124.

Poole, T.B., Taylor, V.J., Fernando, S.B.U., Ratnasooriya, W.D., Ratnayeke, A., Lincoln, G.A., Manatunga, A.M.V.R., Mcneilly, A.S., 1997. Social behaviour and breeding physiology of a group of captive Asian elephants. International Zoo Yearbook 35, 297-310.
Abstract: In 1986 at the Pinnawala Elephant Orphanage, Sri Lanka, a study of 4.4 Asian elephants Elephas maximus was carried out to establish the physical and behavioural changes associated with the female oestrous cycle and the male musth cycle. At Pinnawala the elephants are maintained in mixed-sex groups which has resulted in successful breeding. This paper describes the management of elephants at the orphanage and details their social and reproductive behaviour. It is suggested that successive matings throughout the oestrous cycle and on multiple cycles may be required to ensure successful breeding.

Rasmussen, L.E., Lee, T.D., Zhang, A., Roelofs, W.L., Daves, G.D.Jr., 1997. Purification, identification, concentration and bioactivity of (Z)-7-dodecen-1-yl acetate: sex pheromone of the female Asian elephant, Elephas maximus. Chemical Senses 22, 417-437.
Abstract: In their natural ecosystems, adult male and female Asian elephants, Elephas maximus, live separately. For several weeks prior to ovulation, female elephants release a substance in their urine which elicits a high frequency of non-habituating chemosensory responses, especially flehmen responses, from male elephants. These responses occur prior to, and are an integral part of, mating. Using bioassay-guided fractionation, quantitatively dependent on these chemosensory responses, a specific sex pheromone was isolated and purified by an alternating series of organic and/or aqueous extractions, column chromatography, gas chromatography and high-performance liquid chromatography. Using primarily 1H-proton nuclear magnetic resonance (NMR) spectrometry and gas chromatography-mass spectrometry (GC-MS) of the urine-derived pheromone and its dimethyl disulfide derivative, we determined the structure of the active compound to be (Z)-7-dodecen-1-yl acetate (Z7-12:Ac). Concentrations of Z7-12:Ac in the female urine increased from non-detectable during the luteal phase to 0.48 microgram/ml (0.002 mM) early in the follicular phase and to 33.0 micrograms/ml (0.146 mM) just prior to ovulation. Bioassays with commercially available authentic synthetic Z7-12:Ac, using 10 Asian male elephants at several locations in the US, demonstrated quantitatively elevated chemosensory responses that were robust during successive tests, and several mating-associated behaviors. Bioassays with Z7-12:Ac with adult male elephants dwelling in more natural social situations in forest camps in Myanmar revealed some differing contextual pre-mating behavioral components. The remarkable convergent evolution of this compound suggests that compounds identified in mammalian exudates that are also present in pheromone blends of insects should be re-evaluated as potential mammalian chemosignals.

Rasmussen, L.E.L., Lee, T.D., Zhang, A., Roelofs, W.L., Daves, G.D., 1997. Purification, Identification, Concentration and Bioactivity of (Z)-7-Dodecen-1-yl Acetate: Sex Pheromone of the Female Asian Elephant, Elephas maximus. Chemical Senses 22, 417-437.
Abstract: In their natural ecosystems, adult male and female Asian elephants, Elephas maximus, live separately. For several weeks prior to ovulation, female elephants release a substance in their urine which elicits a high frequency of non-habituating chemosensory responses, especially flehmen responses, from male elephants. These responses occur prior to, and are an integral part of, mating. Using bioassay-guided fractionation, quantitatively dependent on these chemosensory responses, a specific sex pheromone was isolated and purified by an alternating series of organic and/or aqueous extractions, column chromatography, gas chromatography and high-performance liquid chromatography. Using primarily ¹H-proton nuclear magnetic resonance (NMR) spectrometry and gas chromatography-mass spectrometry (GC-MS) of the urine-derived pheromone and its dimethyl disulfide derivative, we determined the structure of the active compound to be (Z)-7-dodecen-1-yl acetate (Z7-12:Ac). Concentrations of Z7-12:Ac in the female urine increased from non-detectable during the luteal phase to 0.48 ug/ml (0.002 mM) early in the follicular phase and to 33.0 ug/ml (0.146mM) just prior to ovulation. Bioassays with commercially available authentic synthetic Z7-12:Ac, using 10 Asian male elephants at several locations in the US, demonstrated quantitatively elevated chemosensory responses that were robust during successive tests, and several mating-associated behaviors. Bioassays with Z7-12:Ac with adult male elephants dwelling in more natural social situations in forest camps in Myanmar revealed some differing contextual pre-matching behavioral components. The remarkable convergent evolution of this compound suggests that compounds identified in mammalian exudates that are also present in pheromone blends of insects should be re-evaluated as potential mammalian chemosignals.

Tiedemann, R., 1997. Sexual selection in Asian elephants. Science 278, 1550-1551.

Vinod, T.R., Cheeran, J.V., 1997. Activity time budget of Asian elephants (Elephas maximus L.) in Idukki Wildlife Sanctuary, Kerala, South India. Indian Forester 123, 948-951.

Dublin, H., 1996. Elephants of the Masai Mara, Kenya: seasonal habitat selection and group size patterns. Pachyderm 22, 35.

Friend, T., Bushong, D. Stereotypic behaviour in circus elephants and the effect of "anticipation" of feeding, watering and performing.  Proceedings of the 30th International Congress of the International Society for Applied Ethology  14-17 August, 1996, Guelph, Ontario, Canada.  30. 1996.
Ref Type: Conference Proceeding

Koch, E. Orphan elephants go on the rampage. New Scientist 151, 5. 1996.
Ref Type: Magazine Article

Kurt, F., Schmid, J. A comparison of feeding behaviour and body weight in Asian elephants (Elephas maximus). First International Symposium on Physiology and Ethology of Wild and Zoo Animals.  1996.
Ref Type: Conference Proceeding

Mosley, J. Hand-Rearing a Captive-Born Asian Elephant (Elephas maximus): (I) A Study of Physical Development as a Response to the Rearing Regime, and (ii) Social Interactions. Spooner, N. G. and Sharp, K. The Ninth UK Elephant Workshop.  36-65. 1996. England, The North of England Zoological Society. 1996.
Ref Type: Conference Proceeding

Poole, J., 1996. Coming of Age with Elephants: a Memoir. Hodder and Stoughton, New York.

Rasmussen, L.E.L., Munger, B.L., 1996. The sensorineural specializations of the trunk tip (finger) of the Asian elephant, Elephas maximus. Anatomical Record 246, 127-134.
Abstract: BACKGROUND: The dorsal extension of the tip of the trunk of Asian elephants (Elephas maximus), often referred to as "the finger," possesses remarkable mechanical dexterity and is used for a variety of special behaviors including grasping food and tactile and ultimately chemosensory recognition via the vomeronasal organ. The present study describes a unique sensory innervation of this specialized region of the trunk. METHODS: The tip of the dorsal aspect of the trunk is referred to as the trunk tip finger and has been studied grossly in 13 living elephants. One tip from a male Asian elephant was obtained for histologic study when it was accidentally severed. The tissue was fixed in 10% neutral buffered formalin, and portions were either sectioned frozen or embedded in paraffin and serial sectioned. Sections were stained with silver in both cases. RESULTS: The skin of the trunk tip finger differs from that of the surrounding areas; it contains a high density of free nerve endings, numerous convoluted branched small corpuscles, and vellus vibrissae that resemble vellus hairs, which do not protrude beyond the skin surface. The finger is thus densely innervated with three distinctive types of sensory terminals. Corpuscular receptors consist of small Pacinian corpuscles and convoluted branched simple corpuscles. Both are present in the superficial dermis. Abundant regular vibrissae are present in the skin surrounding the trunk tip finger. Short vibrissae that do not protrude from the skin surface, referred to as vellus vibrissae, are abundant in the finger tip. Both types of vibrissae are innervated by hundreds of axons resembling the mystacial vibrissae of rodents. Free nerve endings are numerous in the superficial dermis, often making intimate contact with the basal cells of rete pegs. CONCLUSIONS: The dorsal finger of the trunk tip of Asian elephants has a unique sensory innervation that resembles aspects of sensory innervation of mystacial skin of rodents or lip tissue of monkeys. This dense sensory innervation can be correlated with the tactile ability of these animals to use the trunk finger to grasp small objects for feeding and to insert chemically active samples into the ductal orifices of the vomeronasal organ for subsequent chemosensory processing.

Rasmussen, L.E.L., Hall-Martin, A.J., Hess, D.L., 1996. Chemical profiles of African bull elephants, Loxodonta africana: physiological and ecological implications. Journal of Mammalogy 77, 422-439.
Abstract: This study reports concentrations of testosterone and dihydrotestosterone in both serum and temporal-gland secretion of male African elephant (Loxodonta africana), including radiocollared elephants, and identifies a spectrum of volatile components in the temporal-gland secretions. Androgens in the serum (testosterone and dihydrotestosterone) were measured in 111 adult male African elephants, ages 21-40 years, from two national parks in South Africa during several  years and seasons.  About one-fifth (18.6%) of these mature, male, African elephants exhibited dramatically increased concentrations of testosterone in serum characteristic of male Asian elephants during musth.  In Krueger National Park, six radiocollared male African elephants, ages 25-35 years, were tracked and serially samples for both serum and temporal-gland secretions during a 5-year period. Concentrations of testosterone in serum and temporal gland secretions were elevated cyclically at times when typical musth behaviors, including aggression, were observed.  This study reports the first chemical characterization of the volatile compounds of the temporal gland secretions from male African elephants in musth. It reveals many similarities between the chemical constituents of the temporal-gland secretions of these male African elephants and the compounds identified in male Asian elephants.  In addition, several compounds, not previously identified in temporal-gland secretions of African elephants, are described.  Such chemical data support the behavioral observations by ourselves and other researchers that male African elephants experience musth. Especially convincing are the concurrent hormonal and chemical data from the radiocollared males during episodic periods of behavioral musth. Implications of the incidence of musth in the past and present ecology of African elephants are discussed in view of the increasing compression within national parks.

Schmid, J., Kurt, F. Stereotypes in captive Asian elephants (Elephas maximus). First International Symposium on Physiology and Ethology of Wild and Zoo Animals.  1996.
Ref Type: Conference Proceeding

Thakuria, D.B., Barthakur, T., 1996. Management of musth in a male African elephant by chemical sedatives in the Assam state zoo, Guwahati. Indian Veterinary Journal 73, 339-340.

Turkalo, A., 1996. Studying forest elephants by direct observation in the Dzanga clearing: an update. Pachyderm 22, 60.

Carney, E.L., 1995. The greatest show on earth. Iowa State University Veterinarian 57, 53-54.

Desai, A.A., Johnsingh, A.J.T. Social Organization and Reproductive Strategy of the Male Asian Elephant (Elephas maximus).  532. 1995. Bombay, India, Bombay Natural History Society; Oxford University Press.
Ref Type: Abstract

Desai, A.A., Johnsingh, A.J.T., Joshua, J. Techniques for the Study of Ranging Behaviour of the Asian Elephant (Elephas maximus).  532-533. 1995. Bombay, India, Bombay Natural History Society; Oxford University Press.
Ref Type: Abstract

Fowler, M.E., 1995. Restraint and handling of wild and domestic animals. Iowa State University Press, Ames, Iowa, USA.

Kangwana, K., 1995. Avoidance of Maasai by African Elephants (Loxodonta africana). In: Daniel, J.C. (Ed.), A Week with Elephants; Proceedings of the International Seminar on Asian Elephants. Bombay Natural History Society; Oxford University Press, Bombay, India, pp. 529-530.

Masson, J.M., McCarthy, S., 1995. When elephants weep : the emotional lives of animals. Delacorte, New York.

Rasmussen, L.E.L., 1995. Evidence for long-term chemical memory in elephants. Chemical Senses 20, 762.
Abstract: Asian elephants may have a mechanism to avoid close inbreeding that is mediated through the chemical senses. During the year-long postnatal, nursing period and for a number of years afterwards, young elephants live in intimate association with their mothers and closely related females. Although receptor cells were not detected in the vomeronasal organ (VNO) of newborn elephants (Rasmussen et al., 1993. Chemical Senses 18:618), the VNO is presumed to mature within 4-17 weeks postnatally to coincide with the first recognizable flehmen responses by young elephants. Over the past decade we have recorded the chemosensory-oriented behaviors of five calves during their first year. These young elephant, living within their natal group, were observed to flehmen to the urine from their mother twice as frequently as to urine from other females, both relatives and non-relatives. In the wild, by gradual exclusion, and in captivity, by actual removal for management reasons, prepubertal males are physically separated from their natal group. As adults, these males, prior to mating, cue into pheromones in preovulatory urine with a high frequency of flehmen responses (Rasmussen et al., 1993. J. Chem. Ecol. 19:2115). However, individual cues in maternal urine may override estrous cues. Adult males did not respond with high frequency to estrous urine from their mothers. However, chemical extracts of estrous maternal urine elicited high responses by male offspring, suggesting that the individual identity cues have been removed. We hypothesized that young elephant calves imprint on maternal urine and that they retain a chemical memory of this maternal urine over years. We have tested elephants who have been physically distant from their mothers for two to twenty-seven years. These offspring demonstrated a significantly higher response to maternal urine, whether recently collected or stored frozen since the test elephant's postnatal period, than to all other controls including long-time-ago-familiar, unrelated or non-maternally related urine, recently familiar urine, and non-maternal, lactating urine. Our data suggest chemical memory, via maternal urine, may allow filial-to-maternal recognition over time and space separations.

Turkalo, A., Fay, J.M., 1995. Studying forest elephants by direct observation. Pachyderm 20, 45-54.

Black, J., Brockett, R.C., Makowitz, T. A nocturnal behavioral study of three African elephants (Loxodonta africana africana) at Zoo Atlanta. Proceedings of the 20th National Conference of the American Association of  Zoo Keepers.  10-11. 1994. Atlanta, Georgia, October 10-14, 1993.
Ref Type: Conference Proceeding

Hart, B.L., Hart, L.A., 1994. Fly switching by Asian elephants: tool use to control parasites. Animal Behaviour 48, 35-45.
Abstract: A type of tool used by Asian elephants (Elephas maximus) alluded to over a century ago by C. Darwin is their use of branches as a switch, allegedly to repel flies. In a study of Asian elephants used to take tourists for rides into a national Park in Nepal the elephants were observed to grab branches spontaneously and switch with them during the rides and to switch frequently with branches presented to them at the stable. A study of 15 adult working female elephants was conducted to determine whether the use of switches was related to intensity of fly harassment, and if the behaviour functioned to reduce fly intensity around the elephants. Elephants were presented with switches resembling those they spontaneously picked up during rides. At daybreak (06.00 h), when no flies were present, the median switching rate was about 30 per 10 min. At 4 other observation times during the day (approximately 08.00, 11.00, 15.00 and 18.00 h), when fly counts ranged from a median of 1.5 to 4.0 flies on and around the elephants, the median switching rate ranged from 150 to 186 per 10 min. A comparison of switching rate with daytime temperature changes and feeding schedules indicated that switching is not a manifestation of confinement stereotypy or a behaviour related to cooling the body. In an experiment on 8 elephants conducted during the time of day when fly activity was most intense (approximately 11.00 h), fly counts were conducted for 10 min when no branch was available and immediately thereafter for 10 min when the elephants were presented with branches. The median fly count was significantly reduced by 43%. Elephants of the study sometimes modified the branches by removing side stems or shortening the branch. The frequency of fly switching by wild elephants is not currently known. However, among the captive elephants of this study, fly switching would appear to be one of the most frequently employed instances of tool use. [See also B.L. Hart (1990) Neuroscience Biobehavioral Reviews, 14: 273-294; S. Chevalier-Skolnikoff & J. Liska (1993) Animal Behaviour, 46: 209-219].

Hart, L.A., 1994. The Asian elephants-driver partnership: the drivers'perspective. Applied Animal Behaviour Science 40, 297-312.

Huang, H.B., Wellner, D., Naude, R., Oelofsen, W., Oosthuizen, M.M., Breslow, E., 1994. Amino acid sequence and properties of vasopressin-associated elephant neurophysin. Int J Pept Protein Res 44, 270-277.
Abstract: The primary structure of an elephant neurophysin, homologous to vasopressin-associated neurophysins, is reported. The protein contains a Tyr for Asn substitution at position 75, a position in direct contact with residues 77 and 78 of the monomer-monomer interface. This Tyr residue therefore serves as a potential reporter of the path involved in the long-range linkage between peptide binding and dimerization in this system. NMR studies of the protein in unliganded and liganded states demonstrated normal dimerization properties and the expected increase in dimerization associated with binding peptide. In keeping with an elevated pKa of 11.1 assigned to Tyr-75 by UV spectrophotometric titration, the NMR signals from the 3,5 and 2,6 ring protons of Tyr-75 were shifted 0.3 and 0.2 ppm upfield, respectively, relative to their positions in small peptides, indicating significant shielding and/or hydrogen bonding. The Tyr-75 ring proton signals narrowed slightly, with no discernible change in chemical shift, on conversion from dimer to monomer in the unliganded state. Ring protons of Tyr-49, distant from the monomer-monomer interface, but adjacent to the peptide-binding site, were markedly perturbed by dimerization, in accord with their behavior in bovine neurophysins. The results suggest that the secondary and tertiary structure of the region 75-78 is largely unchanged by dimerization, and argue against an important role for this region in dimerization-mediated conformational changes that alter the binding site in the unliganded state.(abstract truncated at 250 words).

Jayewardene, J., 1994. The Elephant in Sri Lanka. WHT Publications Ltd., Colombo, Sri Lanka.

Maguire, R. Standardized Training and Reinforcement System.  1-34. 1994.  Richard "Army" Maguire.
Ref Type: Report

Mikota, S.K., Sargent, E.L., Ranglack, G.S., 1994. Medical Management of the Elephant. Indira Publishing House, West Bloomfield MI.

Poole, J.H., 1994. Sex differences in the behaviour of African elephants. In: Short, R.V., Balaban, E. (Eds.), The Differences Between the Sexes. Cambridge University Press, Cambridge, pp. 331-346.

Redmond, I. Status and environmental enrichment for captive elephants. Proceedings of the 7th UK Elephant workshop.  71-100. 1994.
Ref Type: Conference Proceeding

Roocroft, A., Zoll, D.A., 1994. Managing Elephants:An Introduction to Their Management and Training. Fever Tree Press, Ramona, CA.

Savage, A., Rice, J.M., Brangan, J.M., Martini, D.P., Pugh, J.A., Miller, C.D., 1994. Performance of African elephants (Loxodonta africana) and California Sea Lions (Zalophus californianus) on a two-choice object discrimination task. Zoo Biology 13, 69-75.

Schmid, J., Zeeb, K., 1994. The establishment of the paddocks for keeping elephants in the circus. Deutsche Tierarztliche Wochenschrift 101, 50-52.
Abstract: The guidelines for keeping, training and using animals in circuses and similar institutions, which are made in connection with the law for prevention of cruelty to animals, claim to keep elephants daily 1 hour unshackled in a group in a paddock. The effect of the paddock on social, play behaviour, and the stereotypic movements of circus elephants is discussed. Parameters for housing and managing captured elephants are based on observations of their normal behaviour in nature. A pilot study with 29 elephants in 4 circuses showed that the paddock enabled the elephants to carry out social and comfort behaviour more frequently than when shackled. The stereotypic movements were nearly absent by keeping the elephants in the paddock.

Sukumar, R., 1994. Elephant Days and Nights: Ten Years with the Indian Elephant. Oxford University Press, New Delhi.

Chevalier, S., Liska, J., 1993. Tool use by wild and captive elephants. Animal Behavior 46(pt2), 209-219.
Abstract: This study describes the tool-use potentials of captive African and Asian elephants, Loxodonta africana and Elephus maximus, respectively, and wild African elephants and explores the ecological and evolutionary significance of this behaviour. Comparison of the types, diversity and frequency of tool use by captive and wild elephants with other known tool-using mammals is offered as a means for evaluating the evolutionary significance of tool-using behaviour. Several hypotheses concerning the origin and evolutionary significance of tool use are examined, and it is proposed that in elephants, tool use may serve as adaptations enabling these furless, large-bodied tropical land mammals to cope with ectoparasites and thermoregulation.

Cupane, A., Leone, M., Vitrano, E., Cordone, L., Hiltpold, U.R., Winterhalter, K.H., Yu, W., DiIorio, E.E., 1993. Structure-dynamics-function relationships in Asian elephant (Elephas maximus)myoglobin. An optical spectroscopy and flash photolysis study on functionally important motions. Biophys J 65, 2461-2472.
Abstract: In this work we report the thermal behavior (10-300 K) of the Soret band lineshape of deoxy and carbonmonoxy derivatives of Asian elephant (Elephas maximus) and horse myoglobins together with their carbon monoxide recombination kinetics after flash photolysis; the results are compared to analogous data relative to sperm whale myoglobin. The Soret band profile is modeled as a Voigt function that accounts for the coupling with high and low frequency vibrational modes, while inhomogeneous broadening is taken into account with suitable distributions of purely electronic transition frequencies. This analysis makes it possible to isolate the various contributions to the overall lineshape that; in turn, give information on structural and dynamic properties of the systems studied. The optical spectroscopy data point out sizable differences between elephant myoglobin on one hand and horse and sperm whale myoglobins on the other. These differences, more pronounced in deoxy derivatives, involve both the structure and dynamics of the heme pocket; in particular, elephant myoglobin appears to be characterized by larger anharmonic contributions to soft modes than the other two proteins. Flash photolysis data are analyzed as sums of kinetic processes with temperature-dependent fractional amplitudes, characterized by discrete pre-exponentials and either discrete or distributed activation enthalpies. In the whole temperature range investigated the behavior of elephant myoglobin appears to be more complex than that of horse and sperm whale myoglobins, which is in agreement with the increased anharmonic contributions to soft modes found in the former protein. Thus, to satisfactorily fit the time courses for CO recombination to elephant myoglobin five distinct processes are needed, only one of which is populated over the whole temperature range investigated. The remarkable convergence and complementarity between optical spectroscopy and flash photolysis data confirms the utility of combining these two experimental techniques in order to gain new and deeper insights into the functional relevance of protein fluctuations.

Diephuis, E.P., 1993. Oestrus and pregnancy detection by flehmen-like responses of Asian bull elephants to urine samples of Asian female elephants. Zoologische Garten 63, 235-245.
Abstract: Urine samples were taken twice a week over a period of 130 days from 12 female Asian elephants of various reproductive status at 4 Dutch zoos. Pools, each of 12 urine samples from 6 cows, were offered to 4 male Asian elephants at 3 Dutch zoos. Flehmen responses by the bulls to the individual urine samples were recorded to detect oestrus and pregnancy in the cows. Serum progesterone data were used as a reference. During the tests, one of the 4 bulls showed hardly any flehmen responses; the others showed flehmen responses, with considerable variation between the 3 bulls in number of responses per sample (0.68±0.90, 0.93±1.24 and 1.14±1.52 per sample resp.). Correlations of flehmen responses between bulls were very low or moderate and negative. The repeatability of the response to urine samples averaged 64%. In the present study oestrus could not be detected by recording flehmen responses, and the information obtained about pregnancy was not conclusive. Several peaks of flehmen responses scattered over the 130-day period per cow were found, not exclusively during periods of low serum progesterone level (i.e. oestrous periods). Peaks for individual bulls did not usually match those of the other bulls. Urine samples from 2 cows evoked relatively few flehmen responses, which might indicate pregnancy. One of these 2 cows gave birth to a full-term calf on 1 June 1992. Pregnancy of the 2nd cow could not be confirmed. The present study showed that bulls display flehmen responses to urine samples from cows regardless of whether the cow is present or is familiar to the bull. Probably musth did not affect the interest of bulls in urine samples during flehmen tests. However, the maturity of the bull and the degree of sexual activity the bull displays may influence the interest of bulls in urine samples.

Kabigumila, J., 1993. Feeding habits of elephants in Ngorongoro Crater, Tanzania. African Journal of Ecology 31, 156-164.
Abstract: Observations on the feeding habits of elephants were made from June 1984 to May 1985. The elephants' diet comprised at least 36 plant species ranging in size from big trees to small herbs. Elephants ate mostly sedges (Cyperus immensus in particular) and tree browse during the dry season, and legumes (particularly Trifolium masaiense) and grass (mainly Chloris gayana, Cynodon dactylon and Panicum repens) during the wet season. Elephants browsed on and damaged Acacia xanthophloea. Greatest damage occurred to saplings and was mostly done in the dry season when elephants ate significant amounts of this species. It is concluded that since elephant damage was seasonal, most saplings would recover during the wet season.

Koene, P., Jansen, R.G., Nichelmann, M.ed., Wierenga, H.K.ed., Braun, S. Elephant-caretaker bond and show performance in a zoo. Proceedings of the International Congress on Applied Ethology Berlin 1993: 3rd joint meeting.  1993. Darmstadt, Germany, KTBL Kuratorium fur Technik und Bauwesen in der Landwirtschaft.
Ref Type: Conference Proceeding

Rasmussen, L.E.L., Schmidt, M.J. Filial-maternal chemorecognition in Asian elephant. American Society of Mammmalogists . 1993.
Ref Type: Abstract

Rasmussen, L.E.L., Lee, T.D., Daves, G.D., Schmidt, M.J., 1993. Identification of indolo [2,1-b] quinazoline-6,12-dione in the pre-ovulatory, estrous urine of Elephas maximus. Journal of Chemical Ecology 19, 2115-2128.

Rasmussen, L.E.L., Lee, T.D., Daves, G.D.Jr., Schmidt, M.J., 1993. Female-to-male sex pheromones of low volatility in the Asian elephant, Elephas maximus. Journal of Chemical Ecology 19, 2115-2128.
Abstract: In their natural ecosystems, the sexes of Asian elephant, Elephas maximus, live separately.  For several weeks prior to ovulation, the urine and cervical mucus of female Asian elephants contain extractable chemical agents of low volatility that elicit a high frequency of flehmen responses from bull elephants as an integral part of mating.  Subsequent to flehmen responses, male sexual arousal occurs and, if the female is available, mating results.  During the course of our project to determine the agent(s) and describe the responses associated with female to male sexual communication, we have identified an unusual compound.  This compound, apparently the sole component of the active fraction, was identified by mass, proton nuclear magnetic resonance, ultraviolet/visible, and infrared spectrometries as indolo-[2,1-b]quinazoline-6,12-dione (tryptanthrine).  Exhaustive and repetitive bioassays established that pure authentic (synthetic) typtanthrine was not the compound responsible for the bioresponse.   Rather a coeluting minor component, of low volatility, elicited the male bioresponse.

Rietkerk, F.E., Hiddingh, H., Van Dijk, S., 1993. Hand-rearing an Asian elephant Elephas maximus at the Noorder Zoo, Emmen. Iowa State University Veterinarian 32, 244-252.

Shyan, M.R., Dale, R.H.I., Collins, D., Olson, D., Critser, J.K., Noiles, E.E. Preliminary findings on vocal harmony and behavioral relationship in captive African elephants. Proc. Ann. Elephant Workshop. Proceedings of the Annual Elephant Workshop . 1993.
Ref Type: Conference Proceeding

Tchamba, M.N., Seme, P.M., 1993. Diet and feeding behaviour of the forest elephant in the Santchou Reserve, Cameroon. African Journal of Ecology 31, 165-171.
Abstract: This study reports the diet and feeding behaviour of forest elephants in W. Cameroon. Diet consisted primarily of grass (Pennisetum purpureum). Woody material (leaves and stems) made up the smallest proportion of the diet. 22 species of fruit were eaten of which 7 formed a significant component of the diet: Cucurmis manii, Desplatsia subericarpa, Irvingia gabonensis, Parinari excelsa, Strychnos innocuea, Theobroma cacao and Vitex doniana. The greatest variety of fruit was available during the dry season. The frequent presence of crop seeds in the dung piles indicated that elephant incursions into the farms to feed on crops were regular. Elephants removed bark from 8 species of tree but concentrated on Afzelia bipindensis, Coloncoba welwitschii, Bridelia ferruginea and Terminalia superba. Seventeen species were recorded as browsed. Trees in particular were favoured. Elephants fed mainly by grazing or stripping off fruits. Debarking of trees, breaking of the main stems, and uprooting or pushing over were minor feeding activities.

White, L.J.T., Tutin, C.E.G., Fernandez, M., 1993. Group composition and diet of forest elephants, Loxodonta africana cyclotis Matschie 1900, in the Lope Reserve, Gabon. African Journal of Ecology 31, 181-199.
Abstract: Forest elephants were observed opportunistically during 1984-1991 in lowland rain forest in the Lope Reserve, Gabon. Adult females were generally encountered accompanied by one or more offspring, and adult males were generally solitary. Mean group size was 2.8 individuals, and the mean size of a 'family unit' was 3.5 individuals. Associations of more than 8 elephants were exceptional. Group sizes of elephants at Lope were smaller than those recorded for savanna elephants in E. Africa and those of Asian elephants living in Malaysian rain forest. The diet of elephants at Lope was diverse, including a minimum of 307 items. The bulk of the diet in terms of number of species and quantities eaten, came from leaves and bark (70% of all items recorded). Trees represented 73% of the species eaten. In contrast to savanna-living populations, fruit was an important part of the diet. Fruit of at least 72 species was eaten and the remains of at least 1 species of fruit was found in 82% of 311 fresh dung piles searched over a 1-year period.

Aik, S.S., 1992. Preliminary observations on the training of Burmese elephants using xylazine. New Zealand Veterinary Journal 40, 81-84.
Abstract: A traditional elephant training method was chosen to be modified by the use of xylazine as a sedative and muscle relaxant. Three elephant calves with different degrees of tameness were trained using xylazine. The drug was helpful in the training process. Xylazine made restraint of the elephants much easier and safer. During training, repeated doses of xylazine were used to prevent beatings, the wounds worsening and to pacify the elephants. The tamest elephant calf was punished less and took less time to be trained than the others. It is concluded that it is important to play with elephant calves to win their acceptance of man.

Chandrasekharan, K., Radhakrishnan, K., Nair, K.N.M., Prabhakaran, T., 1992. Some observations on musth in captive elephants in Kerala (India). In: Silas, E.G., Nair, M.K., Nirmalan, G. (Eds.), The Asian Elephant: Ecology, Biology, Diseases, Conservation and Management (Proceedings of the National Symposium on the Asian Elephant held at the Kerala Agricultural University, Trichur, India, January 1989). Kerala Agricultural University, Trichur, India, pp. 71-74.

Cheeran, J.V., Chandrasekharan, K., Radhakrishnan, K., 1992. A case of ochlophobia in a tusker. In: Silas, E.G., Nair, M.K., Nirmalan, G. (Eds.), The Asian Elephant: Ecology, Biology, Diseases, Conservation and Management (Proceedings of the National Symposium on the Asian Elephant held at the Kerala Agricultural University, Trichur, India, January 1989). Kerala Agricultural University, Trichur, India, pp. 176.
Abstract: Full text: An adult captive tusker to be used for ceremonial purpose could not tolerate crowd (ochlophobia - fear of the crowd). The animal was put on 2000 mg of chlorpromazine twice daily orally and behaved normally during the entire festival season of 6 months.  The animal again showed symptoms of fear of the crowd when the owner withdrew the drug.  So the animal was put on 100 mg haloperidol twice daily orally.  Thid relieved the symptoms very well but without sedation compared to chlorpromazine hydrochloride.

Dathe, H.H., Kuckelkorn, B., Minnemann, D., 1992. Salivary cortisol assessment for stress detection in the Asian elephant (Elephas maximus): A pilot study. Zoo Biology 11, 285-289.
Abstract: Effects of introducing an unfamiliar female into an Asian elephant herd at Tierpark Berlin were monitored by means of salivary cortisol assessment.  Saliva samples were obtained from a second female for comparative purposes.  The period of familiarization was characterized by an enhanced cortisol level in both animals, with a maximum on the second day after joining.  Cortisol returned to normal on the following day. Manipulations of the keepers caused a transitory increase on two other days.   Possibilities for the use of this noninvasive method of stress monitoring in various management situations are indicated.

Garai, M.E., 1992. Special relationships between female Asian elephants (Elephas maximus) in zoological gardens. Ethology 90, 187-205.

Nair, P.G., Radhakrishnan, K., Chandrasekharan, K., 1992. Mating behaviour of the Asian elephant in captivity. In: Silas, E.G., Nair, M.K., Nirmalan, G. (Eds.), The Asian Elephant: Ecology, Biology, Diseases, Conservation and Management (Proceedings of the National Symposium on the Asian Elephant held at the Kerala Agricultural University, Trichur, India, January 1989). Kerala Agricultural University, Trichur, India, pp. 38-40.

Payne, K.B., Langbauer, W.R., Jr., 1992. Elephant communication. In: Shoshani, J. (Ed.), Elephants: Majestic Creatures of the Wild. Rodale Press, Emmaus, PA. USA, pp. 116-123.

Phillips, P.K., Heath, J.E., 1992. Heat exchange by the pinnae of the African elephant (Loxodonta africana). Comparative Biochemistry and Physiology [A] 101, 693-699.
Abstract: 1. Surface temperatures of the pinnae of four female African elephants were measured at ambient temperatures between 14 and 32 degrees C using infrared thermography. Instantaneous heat losses calculated using those values ranged from 10.67 to 76.2 W under the observed conditions. 2. Using a value of 17 kcal/kg/day, those heat losses account for 0.65-4.64% of the animals' standard metabolic rates, considering one side of one ear only. 3. A model of heat flow across a flat vertical plate was constructed and compared to the actual values. Up to 100% of an African elephant's heat loss needs can be met by movement of its pinnae and by vasodilation. 4. Thermography indicates that the temperature distribution pattern across the pinna changes with ambient temperature and that areas of specialized motor control exist

Ruggiero, R.G., 1992. Seasonal forage utilization by elephants in central Africa. African Journal of Ecology 30, 137-148.
Abstract: Foraging by adult African elephants was studied in the Terminalia/Combretum woodlands of Manovo-Gounda-St Floris National Park, Central African Republic where dominant grasses were Hyparrhenia rufa and Andropogon gayanus. Elephants spent nearly 17 h/d feeding and had a mean daily intake of nearly 7% of liveweight and tended to feed more at night than during the day. Seasonal variation in feeding time, browsing and grazing rates, weight per trunkful, and defaecation were used to estimate forage intake and gross assimilation efficiency. Daily forage consumption was consistent with the range reported in previous literature, but gross assimilation efficiency was lower indicating relatively poor forage quality during the dry season. The grass:browse ratio in the diet was 3.5:1 DW calculated on an annual basis. Combined with increased human disturbance and decreased forage availability during the dry season, elephants experienced nutritional stress resulting in loss of condition. Defaecation rates, which may be useful for estimating elephant numbers or occupance, are discussed and show that extrapolating numbers of forest elephants from data gathered from savanna populations must be treated with caution due to the differential proportions of grass and browse in their diets.

Tobler, I., 1992. Behavioral sleep in the Asian elephant in captivity. Sleep 15, 1-12.
Abstract: Sleeping behavior was investigated during 294 nights for female Asian elephants (circus: n = 7; zoo: n = 5; including an infant). The animals were recorded continuously on time-lapse video tapes for 7-16 days consecutively. Seasonal changes in sleep behavior were studied by comparing summer (16-day) and winter (13-15-day) recordings; and sleep development was assessed by recording a mother and her infant for three consecutive nights per month for 15 months (age 5-19 months). Sleep occurred in a recumbent (RS) and in a standing position (standing sleep: SS). Although signs of paradoxical sleep (PS) were often evident, the exact onset and end of a PS episode could not be determined. Sleep onset occurred after 2100 hours, and sleep increased progressively reaching a maximum between 0100 and 0400 hours. Total sleep time (TST) in the adults comprised 4.0-6.5 hours per night (including 13.8- 130.9 minutes of SS) and did not differ between the two groups. Seasonal differences were present in TST and in the distribution of sleep within the night; more sleep occurred in the winter. The duration of RS episodes in the adults was 72.0 minutes, a value far below the sleep-cycle length of 124 minutes that others have reported for elephants. TST in the infant decreased during the course of the 15-month recording period from 8.1 hours to 5.1 hours. SS occurred for the first time at the age of 9 months

deVilliers, P.A., Pietersen, E.W., Hugo, T.A., Meissner, H.H., Kok, O.B., 1991. Method of sampling food consumption by free-ranging elephant. South African Journal of Wildlife Research 21, 27.
Abstract: A method is proposed to sample food consumption of free-ranging elephant bulls by tracking in private game reserves bordering the Kruger National Park. Plant communities that characterize these reserves are: Combretum apiculatum with various Grewia spp. dominating the shrub stratum; a woodland community dominated by Colophospermum mopane in the tree and shrub stratum; and mixed C. apiculatum-C. mopane woodland where both species occur in the tree as well as the shrub stratum. As a result of overgrazing, the grass stratum in the whole area is in a retrogressive state of succession with Digitaria eriantha dominating on the better drained soil and Panicum maximum occurring on the heavier soils in association with various tree species. Estimation of bite sizes of leaves, bark, branches, roots and grasses, formed the basis of the technique and was sufficiently accurate for general application in the veld. Two- and 3-dimensional measurements of leaves and bark eaten could be determined directly from branches or roots. In other cases, indirect measurements were taken from similar plant structures in the immediate vicinity. Significant correlations were obtained between the measurements and mass of these plant structures. By combining frequency indexation, linear measurements, and sample weighing, a representative subsample can be determined for use in chemical analyses of food plants.

Haynes, G., 1991. Mammoths, mastodonts, and elephants : biology, behavior, and the fossil record. Cambridge University Pres, Cambridge ; New York.

Rasmussen, L.E.L., Lee, T.D. Purification and initial characterization of a pre-ovulatory pheromone from female Asian elephants (Elephas maximus). Chemical Senses 16, 569. 1991.
Ref Type: Abstract
Abstract: Abstract.  Full text.  Urine from female Asian elephants in the pre-ovulatory phase of the ovarian cycle elicits a high frequency of flehmen responses from Asian bulls in a non-habituating manner.  These flehmen responses are an integral part of the mating sequence and suggest the presence of a sex pheromone.  Extraction and partial purification of components with retention of high biological activity was accomplished several years ago [Rasmussen et al. (1982) Science, 217, 159-162].  Subsequently, standard isolation techniques and molecular weight characterization by conventional mass spectrometric methods proved ineffective.  The pheromone was not identifiable by gas chromatography/ mass spectrometry (both electron impact and chemical ionization) nor by solid probe inlet electron impact mass spectrometry.  The pheromone appeared to be a compound of low volatility, of low molecular weight (200-500) and to be thermally labile.  It was not a peptide. The purification was hampered by close association of high concentrations of inactive components, often aromatics, which possessed similar chromatographic properties.  Purified by an empirically determined series of low pressure and high performance liquid chromatography fractionation sequences, guided at each step and in each preparation by high frequency flehmen responses from Asian bull elephants, the active sex pheromone is apparently a single entity.  Recent developments in field desorption mass spectrometric techniques allow molecular weight determinations on several micrograms of thermally labile substances; by this technique the protonated molecular weight was determined to be 249 and a tentative molecular weight of 248 is assigned.  Further information is presented on the physical and chemical properties of the elephant pheromone including its ultraviolet absorption maximum and it nuclear magnetic resonance spectrum.  Supported by NIH grant HD-19219-06.

Wagner, R.A., Bentz, G.H., 1991. An African elephant tusk pulpotomy: a conservative approach. Proceedings American Association of Zoo Veterinarians 1-5.

Weaver, A.Ch. An Illustrated Bibliography of Elephant Behavior and Conservation.  1991.
Ref Type: Unpublished Work
Abstract: Available from Zoological Society of San Diego, PO Box 551, San Diego, CA 92112-0551 USA.

Wilson, J.F., Mahajan, U., Wainwright, S.A., Croner, L.J., 1991. A continuum model of elephant trunks. J Biomech Eng 113,  79-84.
Abstract: A continuum model is presented that relates the trunk parameters of loading, geometry, and muscle structure to the necessary conditions of static equilibrium. Linear theory for stress-strain behavior is used to describe an elephant trunk for an incremental displacement as the animal slowly lifts a weight at the trunk tip. With this analysis and experimental values for the trunk parameters, the apparent trunk stiffness Ea is estimated for the living animal. For an Asian elephant with a maximum compression strain of 33 percent, Ea is of the order of 10(6) N/m2. The continuum model is quite general and may be applied to similar nonskeletal appendages and bodies of other animals.

Chandra, M.S., Ravoof, A., Khan, I.A., 1990. Feeding behavior of domesticated elephants. Myforest 26, 239-242.

Cooper, K.A., Harder, J.D., Clawson, D.H., Fredrick, D.L., Lodge, G.A., Peachey, H.C., Spellmire, T.J., Winstel, D.P., 1990. Serum testosterone and musth in captive male African and Asian elephants. Zoo Biology 9, 297-306.
Abstract: Testosterone concentrations in serum samples collected weekly over a 5-year period from a young adult male Asian elephant (Elephas maximus) and a young adult male African forest elephant (Loxodonta africana cyclotis) were measured by radioimmunoassay.  Testosterone profiles during this maturational period were compared between the two species and related to the occurrence of musth, a recurring physiological and behavioral condition exhibited by most mature Asian, and some African, bull elephants.  Musth is characterized by secretion from the bull's temporal glands, dribbling urine, and increased aggression.  Serum testosterone concentrations in the Asian bull were elevated substantially between April and September each year, coincident with the presence of temporal gland secretion, urine dribbling, and aggressive behavior. Testosterone levels from April through September averaged (± SEM) 41.2 ± 2.8 ng/ml, compared to 7.6 ± 1.0 ng/ml during the rest of the year.  In contrast, the testosterone profile of the African bull showed greater variability and lower levels overall, the only pattern being a tendency for levels to be lowest from November to February (avg. 6.8 ± 1.5 vs. 10.3 ± 0.8 ng/ml during the rest of the year).  Temporal gland secretion and other signs of musth were first observed in this bull in 1988, at age 17.  While his testosterone values did not show a pattern comparable to that in the Asian bull, average testosterone values were significantly greater in 1988 compared to previous years.  The Asian bull showed sexual attention to preovulatory (estrous) cows whether in musth or not, and exposure to estrous cows did not appear to alter the highly consistent, annual pattern of musth as evidenced by temporal gland flow.

Dhungel, S., Brawner, C., Yoder, J., 1990. Elephant training and management in Nepal. Tigerpaper 17, 1-6.

Gilbert, B. Once a malcontent, Ruby has taken up brush and palette. Smithsonian [December], 40-51. 1990.
Ref Type: Magazine Article

Kiley-Worthington, M. Are elephants in zoos and circuses distressed? Applied Animal Behaviour Science 26[3], 299. 1990.
Ref Type: Abstract

Rasmussen, L.E.L., Hess, D.L., Hall-Martin, A. Chemical profiles of temporal gland secretions from captive Asian bull elephants during musth and from African bull elephants living in wild but crowded conditions. Chemical Senses 15, 628. 1990.
Ref Type: Abstract
Abstract: Full-text: This study compares the volatile components of the temporal gland secretions of captive Asian bull elephants in musth and a distinctive group of wild African bull elephants, confined to a national park.  The captive Asian population has been well studied (Rasmussen et al., 1984; Rasmussen, 1988). Serum testosterone was elevated at specific times; aggressive behaviors occurred concomitantly with temporal gland secretions, although aggression and elevated serum testosterone were not always related.  Selected volatiles among the 23 compounds identified demonstrated concentration changes during the progression of musth, at times simultaneously with alterations in testosterone levels (Rasmussen et al., in press).  The African bull elephants have been monitored, behaviorally and physiologically, by radiocontrolled tracking and monthly sampling during the past 5 years.  Aggressive behaviors similar to those of Asian bull elephants have been documented; serum and temporal gland testosterone were elevated concomitantly in a cyclical fashion similar to musth in Asian elephants.  Chemical characterization of the volatiles of the temporal gland secretions from these bulls revealed several similarities to the compounds described in Asian bulls, including several compounds not previously described in African temporal gland secretions.  It is suggested that these chemicals, or other, more ephemeral compounds, may chemically inform other bulls and cows of the musth-like state of these bulls.

Elephant.  1989. Washington, D.C.], National Geographic special; Distributed by Vestron Video; DNAL Videocassette-no.1154.
Ref Type: Online Source
Abstract: The video is an in-depth look at the elephant and its precarious future.  Covers Asian and African elephants, their behavior, their methods of communication, and their society.

Khit, A. Training working elephants in Laos. Proc.Ann.Elephant Workshop 10.  15-28. 1989.
Ref Type: Conference Proceeding

Kuntze, A., 1989. Dermatopathies in elephants and their treatment. Kleintierpraxis 34, 405-415.

Langbauer, W.R., Jr., Payne, K.B., Charif, R.A., Thomas, E.M., 1989. Reponses of captive African elephants to playback of low-frequency calls. Canadian Journal of Zoology 67, 2604-2607.
Abstract:  We conducted a series of playback trials with captive African elephant to identify behaviors that might indicate perception of calls by conspecifics.  Our findings were as follows. (i) The elephants responded clearly to playbacks of prerecorded low-freqency elephant calls.  Responses included, in rough sequence, lifting and stiffening of ears, vocalization, walking or running towards the concealed speaker, clustering in a tight group, and remaining motionless ("freezing"), with occassional scanning movements of the head.  The occurrence of each of these behaviors increased substantially immediately after the playbacks.  (ii) Elephants responded to full-bandwidth playbacks and to playbacks of calls in which most of the above 25 Hz was filtered out, simulating the effect of frequency-dependent attenuation over distance.  (iii) Elephants did not respond to pure-tone control stimulus similar in frequency and intensity to the filtered elephant calls.  Thus, the observed responses to the elephant calls were not merely responses to an unexpected stimulus, but probably indicate recognition of a biologically meaningful signal.

Nair, P.V., 1989. Development of nonsocial behaviour in the Asiatic elephant. Ethology 82, 46-60.
Abstract: The elephant calf, a defended follower is completely dependent on adults till the age of 3 months.  It begins to explore and attempts to feed at 3 to 6  months, and then becomes partially independent with some feeding on its own.  The characteristics of behavior in adults are examined and the development of this pattern in the calf is traced by analysis of duration, transition and clustering of the behavioral elements. Essential activities like suckling, locomotion, and lying down for rest appear soon after birth whereas elements of feeding, grooming, and play appear only at a later stage.  The calf takes the initiative in suckling and its termination, drinking directly by mouth til the age of 6 months.  The first element of feeding appears at about one week in attempts to pick up and hold objects in the trunk.  Co-ordination of limb, trunk, and mouth movement is achieved by about 1 month.  The calf is strong enough to pull out plants by 6 months when independent feeding begins.  In about a year feeding, drinking and dusting patterns are well developed.

Payne, K., 1989. Elephant talk. National Geographic 176, 264-277.

Poole, J., 1989. Mate guarding, reproductive success and female choice in African elephants. Animal Behavior 37, 842-849.
Abstract: Male guarding of females, male mating success and female choice were studied for 8 years among a population of African elephants, Loxodonta africana.  Males were not able to compete successfully for access to oestrous females until approximately 25 years of age.  Males between 25 and 35 years of age obtained mating during early and late oestrous, but rarely in mid-oestrus.  Larger, older males ranked above the younger, smaller males and the number of females guarded by males increased rapidly late in life.  Body size and longevity are considered important factors in determining the lifetime reproductive success of male elephants.  Oestrous females exercised choice by soliciting guarding behavior from musth, but not non-musth males.  Females in mid-oestrus gave loud, very low frequency calls that may attract distant males and incite male-male competition.  The behavior of oestrous females resulted in their mating with males who were old, vigorous and healthy.

Poole, J.H., 1989. Announcing intent: the aggressive state of musth in African elephants. Animal Behavior 37, 140-152.
Abstract: Predictions derived from game theory suggest that animals should not signal their intentions during conflict situations.  However, during the period of musth, male elephants, Loxodonta africana, announce a state of heightened aggression with signals that are unbluffable.  Since smaller musth males in poor condition are able to dominate larger, normally higher-ranking, non-musth males in good condition, musth provides a useful system with which to examine the possibility of honest signaling of motivation, rather than of fighting ability.  Despite the highly aggressive state of males in musth, escalated contests are extremely rare.  The behaviour of musth and non-musth males suggests that opponents are able to estimate their often rapidly changing roles in the asymmetries with relative accuracy.  Since, unlike most other rutting animals, elephants have asynchronous sexually active periods, resource value varies both with age and the fluctuating sexual state of a particular individual.  It is suggested that musth may be a case where information about resource value is conveyed.

Poole, J.H., Moss, C.J. Elephant mate searching: group dynamics and vocal and olfactory communication. Symposia of the Zoological Society of London.  111-125. 1989.
Ref Type: Conference Proceeding

Povinelli, D.J., 1989. Failure to find self-recognition in Asian elephants (Elephas maximus) in contrast to their use of mirror cues to discover hidden food. J Comp Psych 103, 122-131.

Stout, T. Is elephant training a relic from the past -- can it exist in today's environment? Proc.Ann.Elephant Workshop 10.  29-32. 1989.
Ref Type: Conference Proceeding

Sukumar, R., 1989. Crop raiding by elephants. The Asian Elephant: Ecology and Management. Cambridge University Press, Cambridge, pp. 108-133.

Sukumar, R., 1989. The main study area and study methods. The Asian Elephant: Ecology and Management. Cambridge University Press, Cambridge, pp. 39-59.

Sukumar, R., 1989. Movement and habitat utilization. The Asian Elephant: Ecology and Management. Cambridge University Press, Cambridge, pp. 60-68.

Sukumar, R., 1989. Feeding and nutrition. The Asian Elephant: Ecology and Management. Cambridge University Press, Cambridge, pp. 69-85.

Hromadka, J. Observations made on an Asian elephant (Elephas maximus) during gestation. Proc.Ann.Elephant Workshop 9.  93-105. 1988.
Ref Type: Conference Proceeding

Landres, L. Death among captive elephants: Behavioral observations and management considerations. Proc.Ann.Elephant Workshop 9.  81-89. 1988.
Ref Type: Conference Proceeding

Moss, C., 1988. Elephant memories. Fawcett Columbine (Ballantine), New York.

Niemuller-Hare, C., Gray, C., Liptrap, R. A preliminary report on musth in male Asian elephants. Proc.Ann.Elephant Workshop 9.  106-111. 1988.
Ref Type: Conference Proceeding

Poole, J.H., Payne, K., Langbauer, W.R., Moss, C.J., 1988. The social context of some very low frequency calls of African elephants. Behavioral Ecology and Sociobiology 22, 385-392.

Rasmussen, L.E.L., 1988. Chemosensory responses in two species of elephants to constituents of temporal gland secretion and musth urine. Journal of Chemical Ecology 14, 1687-1711.
Abstract: This report discusses three areas of investigation: (1) The chemical components in the temporal gland secretion (TGS) of Asian (Elephas maximus) and African (Loxodonta africana) elephants were characterized by radioimmunoassay (RIA) for testosterone (T) and dihydrotestosterone(DHT) levels and by on-column capillary column gas chromatographic analysis of volatiles.  An inverse relationship between TGS testosterone levels and (E)-farnesol levels was observed. (2) African elephants responded preferentially toward a particular constituent of African elephant (TGS). (3) Urine from Asian bull elephants in musth was partially fractionated by high-performance liquid chromatography.  Specific chromatographic regions elicited dramatic avoidance responses from female African elephants.  These results support the suggestion that the TGS plays multiple chemocommunicative roles.

Sukumar, R., Gadgil, M., 1988. Male-female differences in foraging on crops by Asian elephants. Animal Behaviour 36, 1233-1235.

Berg, J.K., 1987. Developmental behavior of three African elephant calves (Loxodonta africana) in captivity. Zoologische Garten 57, 171-196.

Hall-Martin, A.J., 1987. The role of musth in the reproductive strategy of the African elephant (Loxodonta africana). South African Journal of Science 83, 616-620.
Abstract: Behavioral and endocrinological data on African elephants ( Loxodonta africana) are integrated to provide a hypothesis of the adaptations reflected in the phenomenon of musth in bulls.  Occupation of home ranges, movements, male dominance hierarchies and intra-specific agonistic behavior are reviewed.  Bulls in musth leave their home range, travel far and fast, imitate more contacts with distant breeding herds, show aggression which overrides normal social male hierarchies, probably mate more frequently than non-musth bulls and then return to their home range.  This behaviour is associated with elevated levels of serum testosterone and dihydrotestosterone. Elephants normally show a high degree of fidelity to sexually segregated adjoining home ranges, which results in regular contact between the same bulls and cows.  This breeding strategy is applicable to older, dominant bulls within the locally resident hierarchy.  The musth adaptation is a second strategy, whereby younger, lower ranking bulls (25-35 years) can ensure more contacts with cows and maximize their chances of breeding. Because musth bulls mate far from their normal ranges the strategy promotes gene flow and ensures outbreeding.  In English with Afrikaans summary.

Poole, J., 1987. Elephants in musth, lust. Natural History 96, 46-55.

Poole, J.H., 1987. Rutting behavior in African elephants: the phenomenon of musth. Behavior 102, 283-316.

Rapaport, L., Haight, J., 1987. Some observations regarding allomaternal caretaking among captive Asian elephants (Elephas maximus). Journal of Mammalogy 68, 438-442.

Sukumar, R., Bhattacharya, S.K., Krishnamurthy, R.V., 1987. Carbon isotopic  evidence for different feeding patterns  in  an Asian  elephant population. Current Science 56, 11-14.

 1986. Elephant talk. Science Digest 94, 15.

 1986. Elephant calls humans can't hear. Science News 129, 122.

Berg, J.K., 1986. Report on the behavioral development of three captive born African elephant calves. Elephant 2, 39-41.
Abstract: The San Diego Wild Animal Park in San Pasqual, California, had three successful African elephant (Loxodonta africana) births in 1982; "Tavi" (female) born 29 January, "Margo" (female) born 24 May and "Tsavo" (male) born 22 June (Lash, 1982).  The gestation period for the calves was estimated to be from 21 to 21 3/4 months.  By the end of their first year, each calf had grown over one foot in height to approximately 49 inches.  Although they eat solid foods, the calves continue to nurse regularly at about 1 hour intervals for a duration of 1.5 minutes over the 8-hour period of observation.

Dudley, J.P., 1986. Notes on training captive elephants. Elephant 2, 15-18.

Gorman, M.L., 1986. The secretion of the temporal gland of the African elephant, Loxodonta africana as an elephant repellant. Journal of Tropical Ecology 2, 187-190.

Haight, J., Rutkowski, C. No pain, no gain?  Breaking tradition and the elephant too. AAZPA Reg.Conf.Proc.Western Region.  1-5. 1986.
Ref Type: Conference Proceeding

Kahl, V.A.L. The study of the reproductive cycle of the African elephant (Loxodonta africana) through correlation of behavior and cytology with serum and urinary hormonal patterns.  1986. Nebraska, Omaha, USA, University of Nebraska.
Ref Type: Thesis/Dissertation

Lake, E.K., 1986. Adjustment process of African elephants to a novel situation. Elephant 2, 32-38.
Abstract: The purpose of this study was to describe the adjustment process of African elephants when separated from companions and introduced into a new enclosure.  Two elephants were observed for fourteen sessions in the noval enclosure and for five sessions in their regular exhibit areas.  Each session lasted thirty minutes and was divided into three ten minute periods.  Seven defined behaviors were tallied by two observers simultaneously.  The proportion of behaviors indicative of investigation and stress revealed when an animal had adjusted to a new situation, i.e. after adjustment, investigative behaviors consistently predominated over behaviors indicative of stress. Each elephant adjusted differently to the novel situation.  This study then mearly documents the obvious; elephants should be slowly introduced to and observed in novel environments.

Lee, P.C., 1986. Early social development among African elephant calves. National Geographic Research 2, 388-401.

Moore, D.E., Doyle, C.E., 1986. Elephant training and ride operations, part I: animal health, cost/benefit and philosophy. Elephant 2, 19-31.
Abstract: Results from a survey, conducted by the authors as employees of the Burnett Park Zoo, show that very few captive elephants in zoos (18 in the USA) are trained for ride operations.  Trained elephants are easily accessable for treatments, are less "bored", and overall are healthier than non-trained elephants, which may be manifested in a longer life span.  The benefits derived from a well planned elephant training and ride operation outweigh the costs incurred.

Morris, M.D., 1986. Large scale deception: deceit by captive elephants? In: Mitchell, R.W., Thompson, N.S. (Eds.), Deception perspective on human and nonhuman deceit. University of New York Press, Albany.

Payne, K.B., Langbauer, W.R., Thomas, E.M., 1986. Infrasonic calls of the Asian elephant (Elephas maximus). Behavioral Ecology and Sociobiology 18, 297-301.
Abstract: Calls at frequencies below the range of human hearing were recorded from two groups of captive Asian elephants (Elephas maximus).  Most of the calls ranged in frequency from 14 to 24 Hz, with durations of 10-15 s (Fig. 1).  With the nearest elephant 5 m from the microphone, sound pressure levels were 85 to 90 dB (re 20 microPa).  These calls occurred in a variety of circumstances.  Elephants are the first terrestrial mammals reported to produce infrasound.  These calls might be important in the coordination of behavior in thick vegetation or among separated groups of elephants.

Wemmer, C., Mishra, H., Dinerstein, E., 1985. Unusual use of the trunk for sound production in a captive Asian elephant: a second case. Journal of the Bombay Natural History Society 82, 187.

Gadgill, M., Nair, P.V., 1984. Observations on the social behaviour of free ranging groups of tame Asiatic elephant (Elephas maximus Linn). Proc. Ind. Acad. Sci. Anim. Sci. 93, 225-233.

Krishnamoorthi, R., La Mar, G.N., Mizukami, H., Romero, A., 1984. A ¹H NMR comparison of the met-cyano complexes of elephant and sperm whale myoglobin. Assignment of labile proton resonances in the heme cavity and determination of the distal glutamine orientation from relaxation data. Journal of Biological Chemistry 259, 8826-8831.
Abstract: The met-cyano complex of elephant myoglobin has been investigated by high field 1H NMR spectroscopy, with special emphasis on the use of exchangeable proton resonances in the heme cavity to obtain structural information on the distal glutamine. Analysis of the distance dependence of relaxation rates and the exchange behavior of the four hyperfine shifted labile proton resonances has led to the assignment of the proximal His-F8 ring and peptide NHs and the His-FG3 ring NH and the distal Gln-E7 amide NH. The similar hyperfine shift patterns for both the apparent heme resonances as well as the labile proton peaks of conserved resonances in elephant and sperm whale met-cyano myoglobins support very similar electronic/molecular structures for their heme cavities. The essentially identical dipolar shifts and dipolar relaxation times for the distal Gln-E7 side chain NH and the distal His-E7 ring NH in sperm whale myoglobin indicate that those labile protons occupy the same geometrical position relative to the iron and heme plane. This geometry is consistent with the distal residue hydrogen bonding to the coordinated ligand. The similar rates and identical mechanisms of exchange with bulk water of the labile protons for the three conserved residues in the elephant and sperm whale heme cavity indicate that the dynamic stability of the proximal side of the heme pocket is unaltered upon the substitution (His----Gln). The much slower exchange rate (by greater than 10(4] of the distal NH in elephant relative to sperm whale myoglobin supports the assignment of the resonance to the intrinsically less labile amide side chain

Ramachandran, K.K., 1984. Observations on unusual sexual behaviour in elephants. Journal of the Bombay Natural History Society 81(3), 687.

Rasmussen, L.E., Buss, I.O., Hess, D.L., Schmidt, M.J., 1984. Testosterone and dihydrotestosterone concentrations in elephant serum and temporal gland secretions. Biology of Reproduction 30, 352-362.
Abstract: Serum and termporal gland secretions (TGS) were obtained from mature wild African (Loxodonta africana) and captive Asian (Elephas maximus).  Samples were obtained from five cows and eight bulls culled for management purposes in Kruger National Park, South Africa, and from four females and two males residing at the Washington Park Zoo, Portland, Oregon.  Our purpose was to describe the levels of androgens, testosterone, and dihydrotestosterone, and to correlate these observations with sex, species, and behavioral status.  Male-female differences in serum T were pronounced in the Asian species, whereas male and female concentrations overlapped in the African elephant serum. Serum T concentrations in African females were > than in Asian females.  Serum DHT reflected T levels, except that the striking elevation of testosterone in Asian bulls during musth was not paralleled by = increases in DHT.  A species difference observed among males was higher serum T levels in nonmusth Asian bulls (1.84-5.35ng/ml) compared to levels in African bulls (0.38-0.68ng/ml), except for one dominant African bull (6.64ng/ml).  This single African value was still considerably lower than the serum T values of the Asian males during musth. These musth values were the highest serum androgen concentrations: T was between 19 and 40ng/ml (average 26.1 ng/ml).  The TSG values of T and DHT were much higher than serum levels except in the Asian female.  T/DHT ratios in TGS were more similar than in serum.  One dominant African bull had a T TGS value of 78ng/ml, which was much higher than the rest of the African males or females, but considerably lower than an Asian bull in musth (547ng/ml).  It seems apparent that a change in androgen status as reflected in serum and TGS levels of T and DHT precedes or is concomitant with overt alteration in behavior in the Asian male.  The temporal gland appears to actively concentrate androgens in both African males and females, but in the Asian male the gland secretes only during musth when the greatest concentration of both T and DHT were observed.  The apparent difference in the degree of temporal gland secretory activity between the 2 species suggests a more specific communicative function within the Asian male.

Sharma, R., Krishnamurthy, K.V., 1984. Behavior of a neonate elephant (Elephas maximus). Applied Animal Behaviour Science 13, 157-161.
Abstract: Observations of behavior of one calf from birth to 5 h post partum, born to an Asian elephant (Elephas maximus), show that such calves are born precocious.  It urinated, defecated and stood unassisted by the mother, walked and attempted suckling within an hour of birth.  The precocity in development appears to have an indirect impact on the nature of its approach to humans.  Thus, calves acquiring an "aunt" at birth seldom approach humans voluntarily, in contrast to those developing such associations later on.  It is suggested that gravid cows be kept as isolated as possible, particularly close to term, if the calves are to be used for commercial purposes.

Siegel, R.K., Brodie, M., 1984. Alcohol self-administration by elephants. Bulletin of the Psychonomic Society 22, 49-52.
Abstract: The anecdotal and historical literature describing intoxication in elephants from femented fruit of alcoholic beverages is reviewed.  Seven African elephants readily self-administered 7% unflavored alcohol solutions, and the results included separation from herd groupings and changes in the frequency and/or duration of several behaviors as scored according to a quantitative observational system.  Alcohol decreased feeding, drinking, bathing and exploration for most animals.  Inappropriate behaviors such as lethargy and ataxia increased for all elephants.  Results are discussed in terms of stress-induced drinking and intoxication.

Siegel, R.K., 1984. LSD-induced effects in elephants: comparisons with musth behavior. Bulletin of the Psychonomic Society 22, 53-56.
Abstract: Musth is a condition observed in male Asiatic elephants and is characterized by aggression and temporal gland secretion.  A classic and controversial 1962 study attempted to induce a musth syndrome in an elephant via treatment with LSD. Two elephants in the present study survived dosages of LSD (.003 -.10 mg/kg) and exhibited changes in the frequency or duration of several behaviors as scored according to a quantitative observational system.  LSD increased aggression and inappropriate behaviors such as ataxia.  Results are discussed in terms of musth and drug-induced perceptual-motor dysfunction.

Wright, P.G., 1984. Why do elephants flap their ears? South African Journal of Zoology 19, 266-269.

Baker, C.M.A., Manwell, C., 1983. Man and elephant.  The "dare theory" of domestication and the origin of breeds. Zeit. Tierzucht. Zuchtungscbiol. 100, 55-75.

Barnes, R.F.W., 1983. Elephant behaviour in a semi-arid environment [Ruaha National Park, Tanzania]. African Journal of Ecology 21, 185-196.

Berg, J.K., 1983. Vocalizations and associated behaviors of the African elephant (Loxodonta africana) in captivity. Zeitschrift fur Tierpsychologie 63, 63-78.
Abstract: This analysis presents the physical characteristics of the vocalizations of the African elephant and describes the associated behavioral contexts of the elephant's communicative system.  One male and 8 female African elephants were systematically observed in a relatively large captive environment.  Their repertoire of sounds constitutes 10 distinct vocalizations which are emitted during 11 behavioral categories.  Although all but one of the sounds is emitted in more than one context, many of the sounds are more characteristic of a specific behavior than others.  An interesting finding is that the fundamental frequency of the emitted sound is significantly correlated to the level of excitement of the individual.  In general, sounds with a low fundamental frequency are emitted within the animals are in a low level of excitement and are important in those behaviors which promote group cohesion and the orderly interactions of individuals.  In contrast, the higher fundamental frequency sounds predominate when the animals are in a high level of excitement and are emitted most often during aggressive type behaviors.  The elephants' vocalizations are associated behaviors in captivity are discussed in comparison to those of their free-living relatives and those of other animals showing similarities across some of the sounds and behaviors.

Dmytriw, R., Olson, D., 1983. A discussion of an ankle abnormality in a young African elephant at the Indianapolis Zoo. Animal Keepers' Forum 10, 20-22.

Dublin, H., 1983. Cooperation and reproductive competition among female African elephants. In: Wasser, S.K. (Ed.), Social behavior of female vertebrates. Academic Press, New York, pp. 291-313.

Gogoi, P.C., 1983. Domestication of Asian elephants (Elephas maximus). Tigerpaper 10, 21-26.

Greenberg, J., 1983. Natural highs in natural habitats. Science News 124, 300-301.

Hess, D.L., Schmidt, M.J., Schmidt, A.M., 1983. Reproductive cycle of the Asian elephant (Elephas maximus) in captivity. Biology of Reproduction 28, 767-773.
Abstract: Serum samples were obtained either weekly or daily from six mature elephant cows for periods of 7 to 30 months from April 1979 to November 1981, to rigorously describe the hormonal and behavioral components of the reproductive cycle in the Asian elephant.  The estrous state of each elephant was concurrently monitored through quantification of the frequency of male urine testing behavior (Flehmen-like response) under controlled conditions.  Concentrations of progesterone and estradiol were determined in 1.0 ml of serum by RIA after diethylether extraction and purification on Sephadex LH-20 columns.  the presence of gonadotropic activity was assessed with a rat LH RIA and by an in vitro mouse Leydig cell bioassay.  Fifteen ovarian cycles averaging 16.3+0.4 weeks in length were observed in six females.  The luteal phase was 10.5+0.3 weeks, and 5.1+0.4 weeks separated subsequent luteal periods. Estradiol concentrations were extremely variable, and no distinct preovulatory E2 surge was associated with the onset of P4 secretion.  However, the onset of P4 release occurred during a single 24h period and reached maximum levels within 2 to 3 weeks.  Peaks of immunoreactive LH were observed, although more than 50% of such "surges" were unrelated to increments in P4 secretion, and no evidence of bioactivity was observed in any serum sample.  Pregnancy was indicated by continued secretion of P4 after the 12-week luteal phase, and elevation levels (400-1200 pg/ml) were seen throughout the first year and during the last 6 months of gestation.  Serum concentrations of P4 fell dramatically before parturition, consistent with a role for this steroid in the regulation of gestational length. Male urine testing behavior was clearly cyclical, and maximal interest during the late interluteal period was followed by a rapid decrease with the onset of P4 secretion.  These data provide the first substantive description of the hormonal milieu of the reproductive cycle, concurrent male behavior, and the serum hormonal patterns at the onset and termination of gestation in this endangered species.

Kaufman, R.L. First response to an elephant attack. Proc.Ann.Elephant Workshop 4.  35-38. 1983.
Ref Type: Conference Proceeding

Moss, C.J., 1983. Oestrous behavior and female choice in the African elephant. Behavior 86, 167-196.

Moss, C.J., Poole, J.H., 1983. Relationships and social structure of African elephants. In: Hinde, R. (Ed.), Primate Social Relationships: an integrated approach. Blackwell Scientific PublicationsHinde,R..

Rees, P.S., 1983. Synchronization of defaecation in the African elephant (Loxodonta africana). Journal of Zoology (Lond) 201, 581-585.
Abstract: The ecological significance of defaecation in African elephants (Loxodonta africana) has been discussed from a number of points of view.  Dropping counts have been used to investigate population size and movements (Wing and Buss, 1970; Law, Parker and Johnstone, 1975); the chemical composition of dung has been examined (Dougall, 1963; Weir, 1972);  and dung decomposition and its role in nutrient cycles has been examined (Coe, 1972) and food consumption of wild elephant herds has been calculated from data on gross assimilation efficiency and drug production (Coe, 1972;Rees, 1982).  This paper presents evidence of a synchronization of defaecation in elephants and discusses its possible ecological significance.

Five elephants plunge over cliff. The Ann Arbor News May 15. 1982.
Ref Type: Newspaper

Barnes, R.F.W., 1982. Elephant feeding behavior in Ruaha National Park, Tanzania. African Journal of Ecology 20, 123-136.

Campbell, A. Care and training of newly imported young African elephants. AAZPA Reg.Conf.Proc.  198-199. 1982.
Ref Type: Conference Proceeding

Eltringham, S.K., 1982. Elephants. Blandfort Books,Ltd., United Kingdom.

Markowitz, H., 1982. On elephants forgetting and bathing. Behavioral Enrichment in the Zoo. Van Nostrand Reinhold Company, New York, pp. 86-93.

Markowitz, H., 1982. Reaffirming that animals are smarter than investigators. Behavioral Enrichment in the Zoo. Van Nostrand Reinhold Company, New York, pp. 132.

Meyer-Holzapfel, M., 1982. Experiments with elephants and tigers in the Swiss National Circus [Reaction from word commands in the training of domestic and wild animals]. Zool. Gart. Jena 52, 271-289.

Rasmussen, L.E., Schmidt, M.J., Henneous, R., Groves, D., Daves, G.D.Jr., 1982. Asian bull elephants: flehmen-like responses to extractable components in female elephant estrous urine. Science 217, 159-162.
Abstract: Flehmen-like responses (urine tests) are one of the characteristic behavioral reactions of male Asian elephants (Elephas maximus) to cow elephants in estrus.  Components of the urine of estrous cow elephants were extracted with organic solvents and partially purified by chromatography and shown to evoke Flehmen-like responses when they were presented to adult bulls.

Seitz, A., 1982. Further data on elephant behavior. Zoologische Garten 52, 15-20.

Sever, Z., 1982. An unusual methold of feeding behavior in captive African elephants. Elephant 2, 140-143.
Abstract: An unusual feeding behavior was observed in a small herd of African elephants at the Safari Park, Ramat Gan, Israel. An elephant was observed to insert her trunk into the rectum of other elephants in order to obtain undigested food.  This behavior was studied in relation to the hierarchy of the elephants within the herd.  Half a year of observation led us to conclude that the elephant higher in hierarchical level would insert her trunk into the rectum of elephants lower in the hierarchy.  The reverse was not observed.

Weary elephant found alive in tree. Columbus Dispatch October 15. 1981.
Ref Type: Newspaper

Allaway, J., 1981. The African elephant's drinking problem. Natural History 90, 30-35.

Hiller, R.B. Elephant training: reasons and results. AAZPA Reg.Conf.Proc.  119-122. 1981.
Ref Type: Conference Proceeding

Mellen, J.D., 1981. Environmental enrichment for servals, Indian elephants and Canadian otters Felis serval, Elephas maximus, Lutra canadensis at Washington Park Zoo, Portland. International Zoo Yearbook 21, 196-201.

Adams, J., Berg, J.K., 1980. Behavior of female African elephants in captivity. Applied Animal Ethology 6, 257-276.
Abstract: The purpose of this research was to ascertain the repertoire of behavior of female African elephants (Loxodonta africana) in captivity.  Seven female African elephants were observed for 558 hours and 22 minutes for a period of one year. This paper gives a detailed description of the activities of the elephants maintained in a relatively restricted environment. Twenty one different kinds of behaviors were observed, 14 of which are considered unique to elephants.  The most frequently occurring behavior was the placement of the trunk of one elephant into or near the mouth of another elephant.  The activities are discussed in terms of: (1) social behaviors; (2) individual behaviors; (3) biological behaviors; (4) dominance hierarchy; (5) four factors derived by statistical factor analysis.

Ali, S.M., 1980. Some aspects of behaviour and breeding biology of the Indian wild elephant. Tigerpaper 7, 9.

Eisenberg, J.F., 1980. Ecology and behavior of the Asian elephant. Elephant Supplement 1, 36-56.