Urinary androstenedione and luteinizing hormone concentrations during musth in a mature African elephant

Musth has not been well documented in captive African elephants. A 37-year-old African bull elephant in the Kansas City Zoological Park was observed during periods of behavioral musth and non-musth. Androstenedione and luteinizing hormone (LH) concentrations in urine were measured by radioimmunoassay. Urinary androstenedione and LH levels were significantly higher in musth urine than in non-musth samples. A positive correlation (P > 0.001) existed between urinary LH and androstenedione concentrations. These results indicate that musth can occur in a zoo-maintained African elephant and that urinary androgen levels are elevated during musth, possibly as the result of LH stimulation of testicular steroidogenesis.     

Levels of aloe mortality with and without elephants in the Thicket Biome of South Africa

Studies concerning the influence of African elephants ( Loxodonta africana ) on vegetation have produced contradictory results; some show minimal or no effect while others report significant elephant-induced effects. Elephants are generalist megaherbivores but will selectively feed from preferred plant species. We investigated the mortality of aloe plants (highly preferred food items for elephants) at five sites with elephants (treatment) and five paired sites without elephants (control) in the Eastern Cape Province of South Africa. A significantly higher proportion of aloes were dead at treatment sites and significantly more aloes that had lost their crown (headless) were found at treatment sites compared with controls. We conclude that although the proportions of dead aloes at treatment sites were significantly higher, it remains unclear whether there is a need to be concerned with the potential small-scale extinction of aloes from parts of the Eastern Cape Province. The observed mortality may merely be an artefact of the loss of large herbivores through disease (e.g. rinderpest) and hunting in the past.     

A simple and accurate method to sex savannah, forest and Asian elephants using noninvasive sampling techniques

We report the development of a reliable and efficient method for molecular sexing of all extant elephant taxa. We developed primers that amplify two short Y-specific fragments (SRY1 and AMELY2) and one longer X-specific fragment (PLP1), developed from elephant sequences in one multiplex PCR. All fragments were designed to be short (< 200 basepairs) for use with degraded DNA and to be 50 basepairs apart to optimize visualization on agarose gels or as electropherograms. The multiplex PCR method matched sexes for at least 97.9% of the noninvasive savannah elephant samples and produced the expected female/male banding patterns for 14 African forest and 11 Asian elephant samples. We found this method to be more robust, efficient and less prone to contamination than previously developed sexing methods for elephants.     

Estimating daily walking distance of captive African elephants using an accelerometer

Two central concerns for elephant husbandry and management are whether zoological enclosures are appropriately sized and the degree to which naturalistic exercise and activity are observed in such enclosures. In order to address these issues, accurate data on the daily walking distance of elephants both in situ and ex situ are necessary. We used an accelerometer, a pedometer that measures step count and activity level, to estimate walking distance in African elephants (Loxodonta africana) at the San Diego Zoo's Wild Animal Park. The accelerometer was worn simultaneously with a GPS unit that recorded actual walking distance. Estimates of walking distance were extrapolated from the accelerometer and compared with actual distances determined by GPS data. The accelerometer was found to overestimate step count, and subsequently walking distance, by including false counts of steps. Extrapolating walking distance based upon stride length measurements did not match actual GPS walking distance. However, activity level output from the accelerometer significantly correlated with actual GPS walking distance. In addition, we report that the rate of movement is comparable to that reported in other zoological settings. We provide a linear regression equation that can be utilized by other institutions to estimate daily walking distance of elephants in their collection who are outfitted with accelerometers.     

Serum testosterone and musth in captive male African and Asian elephants

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 variation 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 profile 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 in temporal gland flow.     

Fine‐scale group structure and demography of African savanna elephants recolonizing lands outside protected areas

Aim Dispersal is a critical component of animal ecology that is poorly understood for most species. In particular, savanna elephants (Loxodonta africana) have been studied for decades in national parks across Africa, but little is known about their dispersal into new or unused habitats or their population dynamics in human‐dominated landscapes. We capitalized on a natural dispersal event of savanna elephants recolonizing communal land in southern Kenya to document their demographic characteristics and genetic relationships. Location Rift Valley province of Kenya. Methods We collected faecal samples and used genetic methods to identify individuals, estimate the sex ratio and evaluate the patterns of relatedness within the female groups and male aggregations. We also measured dung bolus circumference to assign age classes to individuals and estimate the age structure. Results We identified 112 individuals with a sex ratio not different from one (1.32:1.00). The age structure was skewed towards younger elephants (71%), suggesting the potential for rapid growth from reproduction. We detected significantly higher kinship levels within female groups (R = 0.124 ± 0.023), suggesting that family groups colonized the site, but found little support for higher‐order genetic relationships among female groups. Males detected together were unrelated (R = 0.003 ± 0.030). Main conclusions Our results suggest that highly social mammals, such as savanna elephants, disperse into unoccupied habitat as family groups and that a young demographic structure and a large number of males might be expected in establishing populations. These findings highlight the potential value of indirect, non‐invasive methods for assessing elephant herd and demographic characteristics when direct observations are difficult.     

Characterization of tetranucleotide microsatellite loci in the African Savannah Elephant (Loxodonta africana africana)

Most African elephant (Loxodonta africana africana) populations are isolated and thus threatened by a loss of genetic diversity. As a consequence, genetic analysis of African elephant populations will play an increasing role in their conservation, and microsatellite loci will be an important tool in these analyses. Previously published sets of polymorphic microsatellites developed for African elephants are all dinucleotide repeats, which are prone to typing error. Here, we characterize 11 tetranucleotide microsatellite loci in the African elephant. All loci were polymorphic in 32 faecal samples and two tissue samples from 33 individual African savannah elephants.     

Social factors influence ovarian acyclicity in captive African elephants (Loxodonta africana)

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. Zoo Biol 28:1–15, 2009. © 2008 Wiley-Liss, Inc.     

Six new polymorphic microsatellite loci isolated and characterized from the African savannah elephant genome

The African savannah elephant (Loxodonta africana) is a ‘keystone’ species that plays a vital role in regulating the dynamics of both plant and animal communities and yet it is endangered and its numbers have been reduced to approximately 500 000 across their entire continental range. Molecular genetic markers are important tools for providing genetic information useful in formulating effective management and conservation strategies for the surviving elephant populations. We describe the isolation and characterization of six new polymorphic microsatellite markers in the African savannah elephant and demonstrate that these loci can be PCR (polymerase chain reaction)‐multiplexed, a desirable attribute that saves costs in large‐scale microsatellite screening.     

Digestibility of timothy hay by African elephants

Two juvenile, female African elephants (Loxodonta africana) were used in summer and winter trials to determine the apparent digestibility of timothy (Phleum pratense) hay. After 12–14 days of dietary adjustment, dry matter intake and fecal excretion were quantitatively measured for 7 days. Dry matter of timothy hay contained 8.6 and 7.7% crude protein, 57.3 and 44.0% acid detergent fiber, and 6.5 and 6.4% ash during the summer and winter trials, respectively. Estimates of apparent digestibility during summer and winter, respectively, were 39 and 35% for dry matter, 43 and 32% for gross energy (GE), 45 and 30% for crude protein (CP), and 36 and 24% for acid detergent fiber (ADF). While GE and CP digestibility estimates tended (P < .09) to be greater in the summer trial, only the digestibility of ADF was different (P < .05) between summer and winter. Dry matter intake was 1.4–1.6% of body weight (BW), providing an average of 144 kcal of digestible energy per kg BW^0.75. This value is similar to that (155 kcal per kg BW^0.75) used for estimating digestible energy requirements for maintenance of light-breed horses.     

Calculation of longevity and life expectancy in captive elephants

The concepts of longevity (longest lived) and life expectancy (typical age at death) are common demographic parameters that provide insight into a population. Defined as the longest lived individual, longevity is easily calculated but is not representative, as only one individual will live to this extreme. Longevity records for North American Asian elephants (Elephas maximus) and African elephants (Loxodonta africana) have not yet been set, as the oldest individuals (77 and 53 years, respectively) are still alive. One Asian elephant lived to 86 years in the Taipei Zoo. This is comparable to the maximum (though not typical) longevity estimated in wild populations. Calculation of life expectancy, however, must use statistics that are appropriate for the data available, the distribution of the data, and the species' biology. Using a simple arithmetic mean to describe the non‐normally distributed age at death for elephant populations underestimates life expectancy. Use of life‐table analysis to estimate median survivorship or survival analysis to estimate average survivorship are more appropriate for the species' biology and the data available, and provide more accurate estimates. Using a life‐table, the median life expectancy for female Asian elephants (Lx=0.50) is 35.9 years in North America and 41.9 years in Europe. Survival analysis estimates of average life expectancy for Asian elephants are 47.6 years in Europe and 44.8 years in North America. Survival analysis estimates for African elephants are less robust due to less data. Currently the African elephant average life expectancy estimate in North America is 33.0 years, but this is likely to increase with more data, as it has over the past 10 years. Zoo Biol 23:365–373, 2004. © 2004 Wiley‐Liss, Inc.     

Short-term gland secretion in the African elephant: tourist disturbance or social interactions as source for secretion?

The temporal gland is an organ found exclusively in elephants, located midway between the eye and the ear. Hypotheses exist about what factors lead to the release of the secretion but so far few studies have taken place in the wild, especially of secretion in relation to behaviour. It has been suggested that the secretion serves in communication or that it is a symptom of stress. These hypotheses were tested by observing free-ranging elephants in Etosha National Park, Namibia. Factors considered were the impact of carnivores, of herds of other herbivores, of vehicles and of other elephants on the amount of secretion in focal elephant families. Neither other mammals nor tourists, recorded in terms of vehicles, seemed to affect secretion, but the secretion increased with the rate at which members of elephant families displaced each other. It is concluded that the secretion is predominantly related to social behaviour and less to factors extrinsic to the elephant herd.     

Home range, association and related aspects of elephants in the eastern Transvaal Lowveld

Home range and association patterns of elephants Loxodonta africana in private nature reserves adjacent to the Kruger National Park, South Africa, were determined over a period of six years by using radio telemetry. The minimum convex polygon (MCP) and harmonic mean (HM) methods were used for data processing. The minimum and maximum range areas required by females were estimated at 115 and 465 km², respectively, whilst those of males varied between 157 and 342 km². Core areas of females comprised on average 10·1% of the range areas as calculated by the MCP method. Range areas between females in the Klaserie and Timbavati Private Nature Reserves differed significantly. A significant difference in range size was also found between the male population (pooled data) and the females of the Klaserie Private Nature Reserve. Range areas increased from the wet to the dry season. Three male elephant categories were established, namely males utilizing the same area during the study period, those who shifted their range areas and individuals with very large range areas which frequented the area on an irregular basis. A loose relationship existed between males. Association between females can be attributed to kinship and not to the overlapping of range areas. Results indicated avoidance behaviour between females in their core areas, which might partially explain the ability of elephants to tolerate high densities. It is concluded that male elephant hunting should be approached with caution.     

The African savannah elephant population in Kidepo Valley National Park, Uganda: changes in size and structure from 1967 to 2000

The age and sex structure of the elephant population in Kidepo Valley National Park were studied using recognition files. Moreover, population trends were reviewed using past studies. From 1967 to 2000, the elephant population varied between 200 and 500 individuals. Of the minimum number of 374 elephants found to use the Park, 352 were individually identified. Seventy‐nine per cent of these were recorded in 29 families ranging from three to 22 animals with a mean group size of ten. Young animals dominated the population (45% aged 0–9.9 years), while 11% were 10–14.9 years of age and only 18% over 25 years of age. Overall, males constituted 45% of the population, but only 32% of the individuals in the 10–14.9‐year age class were males. Individuals ≥15 years of age displayed a skewed sex ratio of 1 : 1.4 in favour of females. The extent of skew was greatest among mature animals (≥25 years of age), which had four times as many females as males. The skewed age structure in the year 2000 caused by poaching and drought, led to a significant difference between the age distribution in 1970 and 2000. Recent counts suggest that the Kidepo elephant population may be increasing.     

The role of salinity and sodicity in the dieback of Acacia xanthophloea in Ngorongoro Caldera, Tanzania

Dieback of Acacia xanthophloea (Benth.) has opened up the once densely forested Lerai area in Ngorongoro Caldera, Tanzania. Soil samples were taken from profiles in the Ngorongoro Conservation Area and Lake Manyara National Park at sites of dieback and at sites with healthy A. xanthophloea trees. Dieback sites had significantly greater electrical conductivity (EC), water‐soluble Na⁺, K⁺, Cl^?, SO and sodium adsorption ratios (SAR) than healthy sites. The following mean values were recorded: EC (179 versus 70 mS m^?1; P < 0.001, Student's t‐test, n = 8 and 10, respectively; 40–60 cm); Na⁺ (99 versus 30 mmol_c kg^?1, P < 0.001, n = 7 and 8 respectively); K⁺ (11 versus 3 mmol_c kg^?1, P < 0.05); Cl^? (36 versus 7 mmol_c kg^?1, P < 0.01); SO (31 versus 5 mmol_c kg^?1, P < 0.01); and SAR (28 versus 8 mmol l^?1/2, P < 0.01). Water‐soluble Na⁺, Cl^? and SO concentrations in the Lerai profiles have probably resulted in toxicity and osmotic stress which contributed to dieback. Accumulation of salts may have occurred because of reduced flow of freshwater through Lerai and/or flow of water from Lake Magadi into Lerai. Forest recovery may be possible if salinity is reduced. Management strategies for reducing salinity have been implemented and included re‐establishing streams that flow through Lerai. Exclusion of elephants (Loxodonta africana) from Lerai is another management strategy presently under consideration.     

Demography of a population with a long history of poaching and the utility of the individual identification technique as a tool for monitoring intermittently studied elephant populations

In many elephant populations in Africa, adverse effects of poaching including altered age and sex structure and long calving intervals continue to negatively impact the rate of recovery. The study reported here characterized the demographic status of the elephant population in Meru National Park, Kenya, so as to determine the size, age and sex structure, and calving interval of the population and to compare its demographic parameters against those from the relatively stable elephant population at Amboseli. Additionally, a demographic study conducted 7 years earlier enabled us to determine the demographic performance of the population and to explicitly test the utility of the individual identification technique in monitoring intermittently studied populations. We found 392 elephants in the park. The proportion of older individuals was lower, and the calving interval was longer than estimates from Amboseli. We recognized 16% of elephants indexed in the previous study. Our results suggest that the individual identification technique may be useful in monitoring demographic changes including poaching in intermittently studied elephant populations. Overall, our results suggest that although the Meru elephant population showed signs of recovery, poaching continues to imperil its recovery.