Elephant Scar Prevalence in the Kasigau Wildlife Corridor, Kenya: Echoes of Human-Elephant Conflict.

Human-elephant conflict (HEC) compromises crop security and threatens elephant conservation. Most commonly, HEC manifests as crop-foraging as elephants modify natural foraging strategies to incorporate crops. Farmers may retaliate by frightening or harming elephants, leaving scars from inflicted wounds. We assessed the prevalence and distribution of scars on the bodies of African savanna elephants (Loxodonta africana) observed in the Kasigau Wildlife Corridor (KWC), part of the Greater Tsavo Ecosystem of Kenya, where conflict is prevalent. We surmised that scars on the body are largely a result of HEC as opposed to scars on the rump or head, which we attributed primarily to elephant-elephant conflict. We hypothesized that: (1) male elephants would have more scars than females; (2) older males would be more likely to have scars than younger males; and (3) most scars would be located on the bodies of elephants. We assessed scars from a photographic catalogue of elephants from the KWC. In line with our hypotheses, male elephants were more likely to have scars than females (32% of males compared to 6% of females); older males had significantly more scars than younger males (61% compared to 24%); and the majority of scars (89%) were located on the body. Scar presence may be useful as an animal-centered indicator to estimate the prevalence and demographic patterns of HEC.

A pachyderm perfume: odour encodes identity and group membership in African elephants.

Group-living animals that live in complex social systems require effective modes of communication to maintain social cohesion, and several acoustic, olfactory and visual signaling systems have been described. Individuals need to discriminate between in- and out-group odour to both avoid inbreeding and to identify recipients for reciprocal behaviour. The presence of a unique group odour, identified in several social mammals, is a proposed mechanism whereby conspecifics can distinguish group from non-group members. African elephants (Loxodonta africana) live in stable, socially complex, multi-female, fission-fusion groups, characterized by female philopatry, male dispersal and linear dominance hierarchies. Elephant social behaviour suggests that individuals use odour to monitor the sex, reproductive status, location, health, identity and social status of conspecifics. To date, it is not clear what fixed or variable information is contained in African elephant secretions, and whether odour encodes kinship or group membership information. Here we use SPME GC-MS generated semiochemical profiles for temporal, buccal and genital secretions for 113 wild African elephants and test their relationship with measures of genetic relatedness. Our results reveal the existence of individual identity odour profiles in African elephants as well as a signature for age encoded in temporal gland and buccal secretions. Olfactory signatures for genetic relatedness were found in labial secretions of adult sisters. While group odour was not correlated with group genetic relatedness, our analysis identified "group membership" as a significant factor explaining chemical differences between social groups. Saturated and short-chain fatty acids (SCFAs), derived from key volatile compounds from bacterial fermentation, were identified in temporal, buccal and genital secretions suggesting that group odour in African elephants may be the result of bacterial elements of the gut microbiome. The frequent affiliative behavior of African elephants is posited as a likely mechanism for bacterial transmission. Our findings favour flexible group-specific bacterial odours, which have already been proposed for other social mammals and present a useful form of olfactory communication that promotes bond group cohesion among non-relatives in fission-fusion mammals.

The Chemical Ecology of Elephants: 21st Century Additions to Our Understanding and Future Outlooks.

Chemical signals are the oldest and most ubiquitous means of mediating intra- and interspecific interactions. The three extant species of elephants, the Asian elephant and the two African species, savanna and forest share sociobiological patterns in which chemical signals play a vital role. Elephants emit secretions and excretions and display behaviors that reveal the importance of odors in their interactions. In this review, we begin with a brief introduction of research in elephant chemical ecology leading up to the 21st century, and then we summarize the body of work that has built upon it and occurred in the last c. 20 years. The 21st century has expanded our understanding on elephant chemical ecology, revealing their use of odors to detect potential threats and make dietary choices. Furthermore, complementary in situ and ex situ studies have allowed the careful observations of captive elephants to be extended to fieldwork involving their wild counterparts. While important advances have been made in the 21st century, further work should investigate the roles of chemical signaling in elephants and how these signals interact with other sensory modalities. All three elephant species are threatened with extinction, and we suggest that chemical ecology can be applied for targeted conservation efforts.

The relationship of dominance, reproductive state and stress in female horses (Equus caballus).

Maintaining a dominant position in a hierarchy requires energetically expensive aggressive displays and physical exertion. Lab based winner-loser studies, often conducted with individuals from non-social species, have shown that subordinates have higher stress hormone levels than dominant individuals. However, in wild studies on cooperative breeders, displays of aggression used to maintain dominance status are associated with elevated stress hormone levels. The effect of reproductive state on dominance and stress has not been addressed within either of these situations. The purpose of this study was to examine physiological stress levels in relation to dominance rank and reproductive state in a non-cooperative breeder and herbivore, the domestic horse. The social interactions and measured faecal glucocorticoids were recorded in pastured, female horses that were either lactating or non-lactating. While faecal glucocorticoid metabolite level did not differ between reproductive state and rank, activity behaviour demonstrated significant differences between reproductive states. Higher energetic requirements of lactation were reflected in significantly longer bouts of eating and significantly less time spent alert and socializing. As non-cooperative breeders, the social nature of horses does not limit their reproduction or resource acquisition based upon rank, and therefore does not fit with the dominance-stress hypothesis or subordinate-stress hypothesis and instead supports a rank-independent stress hypothesis.

Impacts of environmental pressures on the reproductive physiology of subpopulations of black rhinoceros (Diceros bicornis bicornis) in Addo Elephant National Park, South Africa.

Black rhinoceros are an icon for international conservation, yet little is known about their physiology due to their secretive nature. To overcome these challenges, non-invasive methods were used to monitor rhinoceros in two sections of Addo Elephant National Park, South Africa, namely Addo and Nyathi. These sections were separated by a public road, and the numbers of elephants, predators and tourists were higher in Addo. Faecal samples (n = 231) were collected (from July 2007 to November 2010) from known individuals and analysed for progestagen and androgen metabolite (FPM and FAM, respectively) concentrations. As biotic factors could impact reproduction, we predicted that demographics, FPM and FAM would vary between sections and with respect to season (calendar and wet/dry), climate and age of the rhinoceros. Mean FPM concentrations from pregnant females were seven times higher (P < 0.05) than samples from non-pregnant rhinoceros. Positive relationships were found between monthly temperatures and FPM from non-pregnant females (r (2) = 0.25, P = 0.03) and the percentage of calves born (r = 0.609, P = 0.04). Although FAM peaked in the spring, when the majority of calves (40%) were conceived, no seasonal patterns in male androgen concentrations were found with respect to month of conception and parturition. Females in Addo had a longer inter-calving interval and were less likely to be pregnant (P < 0.05) compared with those in Nyathi. The biotic stressors (e.g. predators and more competitors) within Addo section could be affecting the reproductive physiology of the rhinoceros negatively. Enhanced knowledge about how black rhinoceros populations respond to environmental stressors could guide management strategies for improving reproduction.

Ovarian cycle activity varies with respect to age and social status in free-ranging elephants in Addo Elephant National Park, South Africa.

Free-ranging African elephants live in a fission-fusion society, at the centre of which is the matriarch. Matriarchs are generally older females that guide their families to resources and co-ordinate group defense. While much is known about elephant society, knowledge is generally lacking about how age affects the physiology of wild elephants. Investigation of the ovarian activity of free-ranging elephants could provide insight into the reproductive ageing process, with implications for population management. Faecal samples were collected from 46 individuals ranging in age from 14 to 60 years for a 2-year period, and progestagen metabolite analyses were used to examine relationships between social status, age, season, and ovarian activity in female elephants in Addo Elephant National Park, South Africa. Social status was the strongest predictor of faecal progestagen metabolite concentrations in non-pregnant elephants, with grand matriarchs (n = 6) having the lowest values compared with matriarchs (n = 21) and non-matriarch females (n = 19). Likewise, social status and age were the strongest predictors of faecal progestagen metabolite concentrations in pregnant elephants (n = 27). The number of years since a non-pregnant female gave birth to her last calf (post-partum duration) was longer for older females with a higher social status, as well as during the dry season. Our results indicate that social standing and age of elephants are related to reproductive function, and that older females exhibit reductions in ovarian capacity. These results expand our understanding of reproduction and fertility throughout an elephant's lifespan, and the factors that impact gonadal function in free-ranging females. Given that possible over-abundance of elephants in areas such as Addo Elephant National Park is fuelling the debate over how best to manage these populations, knowledge about the reproductive potential of high-ranking females can provide managers with biological data to identify the best candidates for controlling growth through translocation or contraception.

Chemical signals of elephant musth: temporal aspects of microbially-mediated modifications.

Mature male African (Loxodonta africana) and Asian (Elephas maximus) elephants exhibit periodic episodes of musth, a state in which serum androgens are elevated, food intake typically decreases, aggressiveness often increases, and breeding success is enhanced. Urine is a common source of chemical signals in a variety of mammals. Elephants in musth dribble urine almost continuously for lengthy periods, suggesting that the chemicals in their urine may reveal their physiological condition to conspecifics. We investigated the volatile urinary chemicals in captive male elephants using automated solid phase dynamic extraction (SPDE) and gas chromatography-mass spectrometry (GC-MS). We found higher levels of alkan-2-ones, alkan-2-ols, and some aromatic compounds in urine from males in musth than in urine from non-musth males or from females. Levels of ketones and alcohols increased as the urine aged, likely due to microbial metabolism of fatty acids. Protein-derived aromatic metabolites also increased in abundance after urination, likely due to microbial hydrolysis of hydrophilic conjugates. We suggest that microbes may play an important role in timed release of urinary semiochemicals during elephant musth.

Investigating the impact of rank and ovarian activity on the social behavior of captive female African elephants.

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.

Using behavioral observations and keeper questionnaires to assess social relationships among captive female African elephants.

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.

Estrous state influences on investigative, aggressive, and tail flicking behavior in captive female Asian elephants.

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.

Insect pheromones and precursors in female African elephant urine.

Using automated solid-phase dynamic extraction and gas chromatography-mass spectrometry, our search for urinary chemical signals from ovulatory female African elephants (Loxodonta africana) has revealed the bark beetle aggregation pheromones frontalin, exo-brevicomin, and endo-brevicomin, as well as their precursors and the aphid alarm pheromones (E,E)-alpha-farnesene and (E)-beta-farnesene. Enantiomeric ratios for brevicomins have been determined. Prior discovery of common insect/elephant pheromones in Asian elephants, namely, (Z)-7-dodecenyl acetate and frontalin, suggests that the present findings may yield valuable insights into chemical communication among African elephants.