Elephant prolactin: isolation and characterization

Prolactin was isolated from anterior lobes of elephant pituitary glands. It consisted of 199 amino acids with three disulfide bridges and two tryptophan residues as found in prolactin from other species. The sequence of the NH2-terminal 28 amino acids was determined and shown homologous with the ovine hormone. In comparison with ovine prolactin, a marked difference was seen in the methionine content; the elephant hormone possessed only 18-34% lactogenic potency. The conformation of elephant prolactin was examined by zero order, second order and circular dichroism spectroscopy. The alpha helical content was estimated to be about 60%. In comparison with prolactins from other species, the second order spectra of elephant prolactin suggest that the local microenvironment for one or both tryptophan residues is somewhat different.     

Seasonality in elephant dung decay and implications for censusing and population monitoring in south-western Cameroon

Dung decay rates have been used in the past to correct elephant dung counts to estimates of population numbers. These assume that the dung is in a ‘steady state’ where production rates are equal to disappearance rates. The inter‐seasonal variation in decay rates, however, has never been fully investigated for elephant dung. This study, in the Banyang‐Mbo Wildlife Sanctuary in SW Cameroon, investigated whether a single dung decay rate is really appropriate when converting dung counts to population density estimates. A total of 870 fresh elephant dung‐piles were monitored until they had completely decayed, with at least 60 fresh dung piles located each month of the year between July 1994 to June 1995. The elephant dung decay rate obtained was 55.6% to 66.7% lower than those obtained for the rainforests of Gabon. This has implications for using decay rates obtained elsewhere to compute elephant densities from dung counts. Statistically significant differences were found in the mean duration of dung‐piles between different months and different seasons. Environmental variables, with the exception of rainfall, did not aid in predicting the duration of elephant dung‐piles. The model of Plumptre & Harris was used to estimate the standing crop of dung at any particular time of the year. The results show that, if there was no emigration of elephants from the population, then there was only a three‐month period when the dung was in a ‘steady state’. If emigration occurred during the wet season as is thought to occur then there was no period of ‘steady state’.     

Estrogen and LH dynamics during the follicular phase of the estrous cycle in the Asian elephant

Pituitary and corpus luteum hormone patterns throughout the elephant estrous cycle have been well characterized. By contrast, analysis of follicular maturation by measurement of circulating estrogens has been uninformative. This study tested the ability of a urinary estradiol‐3‐glucuronide radioimmunoassay to noninvasively assess follicular development during the nonluteal phase of the elephant estrous cycle, and to determine the relationship between estrogen production and the “double LH surge.” Daily urine and serum samples were collected throughout seven estrous cycles from three Asian elephants, and urine was collected from an additional three females, for a total of 13 cycles. Serum was analyzed for luteinizing hormone (LH), and urine was analyzed for estrogens and progestins. Elephants exhibited a typical LH pattern, with an anovulatory LH (anLH) surge occurring approximately 21 days before the ovulatory LH (ovLH) surge. The urinary estrogen pattern indicated the presence of two follicular waves during the nonluteal phase. The first wave (anovulatory) began 5 days before the anLH surge and reached a maximum concentration the day before the peak. Thereafter, urinary estrogens declined to baseline for 2 weeks before increasing again to peak concentrations on the day of the ovLH surge. Urinary progestins were baseline throughout most of the follicular phase, increasing 2–3 days before the ovLH surge and continuing into the luteal phase. These results support previous ultrasound observations that two waves of follicular growth occur during the nonluteal phase of the elephant estrous cycle. Each wave is associated with an increase in estrogen production that stimulates an LH surge. Thus, in contrast to serum analyses, urinary estrogen monitoring appears to be a reliable method for characterizing follicular activity in the elephant. Zoo Biol 22:443–454, 2003. © 2003 Wiley‐Liss, Inc.     

Social life of captive Asian elephants (Elephas maximus) in Southern India: implications for elephant welfare

Asian elephants in the wild live in complex social societies; in captivity, however, management often occurs in solitary conditions, especially at the temples and private places of India. To investigate the effect of social isolation, this study assessed the social group sizes and the presence of stereotypies among 140 captive Asian elephants managed in 3 captive systems (private, temple, and forest department) in Tamil Nadu, India, between 2003 and 2005. The majority of the facilities in the private (82%) and temple (95%) systems held a single elephant without opportunity for social interaction. The forest department managed the elephants in significantly larger groups than the private and temple systems. Among the 3 systems, the proportion of elephants with stereotypies was the highest in temple (49%) followed by private system (26%) and the forest department facility (6%); this correlates with the social isolation trend observed in the 3 systems and suggests a possible link between social isolation and abnormal elephant behavior separate from other environmental factors. The results of this study indicate it would be of greater benefit to elephant well being to keep the patchily distributed solitary temple and private elephants who are socially compatible and free from contagious diseases in small social groups at "common elephant houses" for socialization.     

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.     

Survey of the reproductive cyclicity status of Asian and African elephants in North America

The Asian and African elephant populations in North America are not self‐sustaining, and reproductive rates remain low. One problem identified from routine progestagen analyses is that some elephant females do not exhibit normal ovarian cycles. To better understand the extent of this problem, the Elephant TAG/SSP conducted a survey to determine the reproductive status of the captive population based on hormone and ultrasound evaluations. The survey response rates for facilities with Asian and African elephants were 81% and 71%, respectively, for the studbook populations, and nearly 100% for the SSP facilities. Of the elephants surveyed, 49% of Asian and 62% of African elephant females were being monitored for ovarian cyclicity via serum or urinary progestagen analyses on a weekly basis. Of these, 14% of Asian and 29% of African elephants either were not cycling at all or exhibited irregular cycles. For both species, ovarian inactivity was more prevalent in the older age categories (>30 years); however, acyclicity was found in all age groups of African elephants. Fewer elephant females (～30%) had been examined by transrectal ultrasound to assess reproductive‐tract integrity, and corresponding hormonal data were available for about three‐quarters of these females. Within this subset, most (～75%) cycling females had normal reproductive‐tract morphologies, whereas at least 70% of noncycling females exhibited some type of ovarian or uterine pathology. In summary, the survey results suggest that ovarian inactivity is a significant reproductive problem for elephants held in zoos, especially African elephants. To increase the fecundity of captive elephants, females should be bred at a young age, before reproductive pathologies occur. However, a significant number of older Asian elephants are still not being reproductively monitored. More significantly, many prime reproductive‐age (10–30 years) African females are not being monitored. This lack of information makes it difficult to determine what factors affect the reproductive health of elephants, and to develop mitigating treatments to reinitiate reproductive cyclicity. Zoo Biol 23:309–321, 2004. © 2004 Wiley‐Liss, Inc.     

Understanding Long-Term Variations in an Elephant Piosphere Effect to Manage Impacts

Surface water availability is a key driver of elephant impacts on biological diversity. Thus, understanding the spatio-temporal variations of these impacts in relation to water is critical to their management. However, elephant piosphere effects (i.e. the radial pattern of attenuating impact) are poorly described, with few long-term quantitative studies. Our understanding is further confounded by the complexity of systems with elephant (i.e. fenced, multiple water points, seasonal water availability, varying population densities) that likely limit the use of conceptual models to predict these impacts. Using 31 years of data on shrub structure in the succulent thickets of the Addo Elephant National Park, South Africa, we tested elephant effects at a single water point. Shrub structure showed a clear sigmoid response with distance from water, declining at both the upper and lower limits of sampling. Adjacent to water, this decline caused a roughly 300-m radial expansion of the grass-dominated habitats that replace shrub communities. Despite the clear relationship between shrub structure and ecological functioning in thicket, the extent of elephant effects varied between these features with distance from water. Moreover, these patterns co-varied with other confounding variables (e.g. the location of neighboring water points), which limits our ability to predict such effects in the absence of long-term data. We predict that elephant have the ability to cause severe transformation in succulent thicket habitats with abundant water supply and elevated elephant numbers. However, these piosphere effects are complex, suggesting that a more integrated understanding of elephant impacts on ecological heterogeneity may be required before water availability is used as a tool to manage impacts. We caution against the establishment of water points in novel succulent thicket habitats, and advocate a significant reduction in water provisioning at our study site, albeit with greater impacts at each water point.     

Isolation and characterization of growth hormone from blue fox pituitary glands

Growth hormone has been purified to homogeneity from blue fox pituitary glands. It has 191 amino acids with two disulfide bridges and a single tryptophan residue. The somatotropin activity is only 8% when compared with the bovine hormone in the receptor-binding assay. From radioimmunoassay data using baboon antisera to porcine or bovine growth hormone, the fox hormone has 14-17% immunoreactivity of bovine or porcine hormone.     

Reproductive cycle of the elephant

The combination of a few factors, including poor captive reproduction, secession of importation from the wild and advances in hormone detection and ultrasonography, has contributed to the current knowledge on the elephant reproductive cycle. Several reproductive features in elephants differ markedly from other mammals. These include the urogenital tract anatomy, length and structure of the reproductive cycle, the formation of multiple corpora lutea and the type and secretion pattern of reproductive hormones. Being 13-18 weeks in length, the elephant estrous cycle is the longest amongst all studied non-seasonal mammals to date. Progesterone increases 1-3 days after ovulation, indicating the start of the luteal phase, which lasts 6-12 weeks. This is followed by a 4- to 6-week follicular phase that is concluded by two, precisely spaced and timed, LH surges. In general, the first, anovulatory LH surge occurs exactly 19-21 days before the second, ovulatory surge. Normally, a single follicle is ovulated. However, beside a corpus luteum (CL) forming on the site of ovulation, multiple accessory CLs can be found on the ovaries. Unlike many other species, the predominant progestagen secreted by luteal tissues is not progesterone, but rather its 5-alpha-reduced metabolites. The currently known aspects of the unique estrous cycle in Asian and African elephants, covering estrous behavior, circulating hormones, ultrasonography and anatomy of the reproductive organs as well as hormonal manipulation treatment possibilities, will be reviewed here.     

Conservation status of Asian elephants: the influence of habitat and governance

Understanding the drivers of Asian elephant (Elephas maximus) abundance and distribution is critical for effective elephant conservation, yet no such analysis exists despite decades of assessments and planning. We explored the influence of habitat- and governance-related drivers on elephant abundance across the 13 Asian elephant range countries. We tested competing statistical models by integrating a binary index of elephant abundance (IEA) derived from expert knowledge with different predictor variables including habitat, human population, socioeconomics, and governance data. We employed logistic regression and model-averaging techniques based on Akaike’s Information Criterion to identify the best-performing subset among our 12 candidate models and used the model-averaged results to predict IEA in other areas in Asia where elephant population status is currently unknown. Forest area was our strongest single predictor variable. The best performing model, however, featured a combination of habitat and governance variables including forest area, level of corruption, proportional mix of forest and agriculture, and total agricultural area. Our predictive model identified five areas with medium–high to high probability to have populations with >150 elephants, which we believe should be surveyed to assess their status. Asian elephants persist in areas that are dominated by forest but also seem to benefit from a mix of agricultural activities. A relatively low level of corruption is also important and we conclude that effective governance is essential for maintaining Asian elephant populations. Asian elephant populations cannot be maintained solely in protected areas but need well-managed, mixed-use landscapes where people and elephants coexist.     

Human settlement drives African elephant (Loxodonta africana) movement in the Sebungwe Region,Zimbabwe

Understanding the key drivers that influence the potential distribution of herbivore species in changing landscapes has been at the centre of enquiry in wildlife science for many decades. This knowledge is particularly important for keystone species like the African elephant (Loxodonta africana) whose population is declining even in conservation areas. The Sebungwe Region is part of the Kavango‐Zambezi Transfrontier Conservation Area and supports ~4,000 elephants. The Sebungwe Region has lost an estimated 76% of its elephant population over the years. This study aimed to determine how the elephant distribution in the Sebungwe Region was affected by human settlement and whether the patches for elephant distribution were large enough for elephant habitation. The prediction of the potential distribution of the elephant was based on presence‐only data modelled through an ensemble algorithm that combined several candidate models to enhance predictive ability. We observed that human settlement drives the potential distribution of elephants in the Sebungwe Region (test AUC = 0.95), and patches from the model were on average <1.5 km². Our results provide initial insights into the key habitat factors that drive distribution of elephants in the Sebungwe landscape. Future conservation of the elephant could benefit from our study through systematic planning of settlements, which might help minimise human interaction with wildlife.     

Molecular Characterization of Pro-Inflammatory Cytokines Interleukin-1β and Interleukin-8 in Asian Elephant (Elephas maximus)

Interleukin (IL)-1β and IL-8 are pro-inflammatory cytokines produced primarily by monocytes and macrophages in response to a variety of microbial and nonmicrobial agents. As yet, no molecular data have been reported for IL-1β and IL-8 of the Asian elephant. In the present study, we have cloned and sequenced the cDNA encoding IL-1β and IL-8 of the Asian elephant. The open reading frame (ORF) of Asian elephant IL-1β is 789 bp in length, encoded a propeptide of 263 amino acid polypeptide. The predicted protein revealed the presence of IL-1 family signature motif and an ICE cut site. Whereas, IL-8 contained 321 bp of open reading frame. Interestingly, the predicted protein sequence of 106 aa, contains an ELR motif immediately upstream of the CQC residues, common in all vertebrate IL-8 molecules. Identity levels of the nucleic acid and deduced amino acid sequences of Asian elephant IL-1β ranged from 68.48 (Squirrel monkey) to 98.57% (African elephant), and 57.78 (Sheep) to 98.47% (African elephant), respectively, whereas that of IL-8 ranged from 72.9% (Human) to 87.8% (African elephant), and 63.2 (human, gorilla, chimpanzee) to 74.5% (African elephant, buffalo), respectively. The phylogenetic analysis based on deduced amino acid sequenced showed that the Asian elephant IL-1β and IL-8 were most closely related to African elephant. Molecular characterization of these two cytokines, IL-1β and IL-8, in Asian elephant provides fundamental information necessary to progress the study of functional immune responses in this animal and gives the potential to use them to manipulate the immune response as recombinant proteins.     

Endocrine and ultrasound evaluation of a non‐cycling African elephant: Identification of an ovarian follicular cyst

The reproductive rate of captive African elephants is low because of logistical difficulties associated with transporting animals for breeding, the danger of maintaining bulls and medical or physiological problems. There also is growing evidence that a significant number of mature female elephants are not experiencing normal estrous cycles. The case described in this report involves the diagnosis and attempted treatment of an ovarian follicular cyst in an African elephant at the Pittsburgh Zoo. On the basis of serum progesterone analysis, the female exhibited regular ovarian cycles from July 1993 through March 1994, but from November 1994 to the present has not shown any evidence of reproductive cyclicity. In April 1996, a large follicular structure was identified on the right ovary using transrectal ultrasound. In an attempt to luteinize the cyst, 500 μg gonadotropin‐releasing hormone (GnRH) was administered intravenously in October 1996, which stimulated a modest increase in serum luteinizing hormone (LH) (approximately twofold over baseline), but no resumption of ovarian activity. The elephant was treated again 5 months later with a higher dose of GnRH (5 mg, i.v.) with the same results. An ultrasound evaluation in July 1997 indicated the structure was still present. In October 1997, the female was given human chorionic gonadotropin (hCG; 10,000 IU, i.m.), which induced estrus and breeding, but no ovulation or luteinization of the cyst, and she remains acyclic. These results suggest that conventional methods developed in other species for treating ovarian follicular cysts may not necessarily be effective in the elephant. It is also important that reproductive age females be monitored via continuous progesterone analysis and occasional reproductive tract ultrasound evaluations to understand better the etiology of ovarian dysfunction so that effective treatments can be developed to induce consistent ovarian activity. Zoo Biol 18:223–232, 1999. © 1999 Wiley‐Liss, Inc.     

Long distance breeding dispersal of a southern elephant seal

Southern elephant seals range extensively during regular foraging excursions. Despite this they are highly philopatric and long range dispersal is rare. At Gough Island, southern Atlantic Ocean, we observed a breeding adult male elephant seal during September 2009, which had been tagged on its natal beach at Marion Island, southern Indian Ocean, in November 1998. The individual was resighted only once on Marion Island, 6 months after tagging. This 3,860 km movement represents dispersal (and likely gene flow) between distinct populations from different elephant seal geographical provinces. Given the polygynous breeding system of this species, the presence of this single male may have a disproportionate genetic effect on the small number of southern elephant seals breeding at Gough Island.     

Past and present status of the southern elephant seal (Mirounga leonina) at Macquarie Island

The southern elephant seal population at Macquarie Island has undergone a serious decline since regular surveys were commenced in 1949. Approximately 2,900 cows and 250 bulls were counted in the isthmus study area during the 1985 breeding season. Comparisons with 20 other counts made in the same area between 1949 and 1984 showed that, although there was considerable fluctuation between the years, the number of seals ashore has dropped at an average rate of 2.1% per year, resulting in a net decrease of approximately 50% for both males and females. This is similar to the ratio of decline of elephant seal populations in the southern Indian Ocean. The census information from all major elephant seal populations was reviewed and it was concluded that there may be a common factor, or group of factors, acting to reduce the Macquarie Island and Indian Ocean populations, while the populations of the South Atlantic seem to be stable. Several potential explanations for these observations are advanced, but it is concluded that a greater knowledge of the ecology of the seals is needed if the declines are to be understood.     

Physiological stress and refuge behavior by African elephants

Physiological stress responses allow individuals to adapt to changes in their status or surroundings, but chronic exposure to stressors could have detrimental effects. Increased stress hormone secretion leads to short-term escape behavior; however, no studies have assessed the potential of longer-term escape behavior, when individuals are in a chronic physiological state. Such refuge behavior is likely to take two forms, where an individual or population restricts its space use patterns spatially (spatial refuge hypothesis), or alters its use of space temporally (temporal refuge hypothesis). We tested the spatial and temporal refuge hypotheses by comparing space use patterns among three African elephant populations maintaining different fecal glucocorticoid metabolite (FGM) concentrations. In support of the spatial refuge hypothesis, the elephant population that maintained elevated FGM concentrations (iSimangaliso) used 20% less of its reserve than did an elephant population with lower FGM concentrations (Pilanesberg) in a reserve of similar size, and 43% less than elephants in the smaller Phinda reserve. We found mixed support for the temporal refuge hypothesis; home range sizes in the iSimangaliso population did not differ by day compared to nighttime, but elephants used areas within their home ranges differently between day and night. Elephants in all three reserves generally selected forest and woodland habitats over grasslands, but elephants in iSimangaliso selected exotic forest plantations over native habitat types. Our findings suggest that chronic stress is associated with restricted space use and altered habitat preferences that resemble a facultative refuge behavioral response. Elephants can maintain elevated FGM levels for ≥ 6 years following translocation, during which they exhibit refuge behavior that is likely a result of human disturbance and habitat conditions. Wildlife managers planning to translocate animals, or to initiate other management activities that could result in chronic stress responses, should consider the potential for, and consequences of, refuge behavior.     

A safer method for studying hormone metabolism in an Asian elephant (Elephas maximus): accelerator mass spectrometry

Noninvasive hormone assays provide a way to determine an animal's health or reproductive status without the need for physical or chemical restraint, both of which create unnecessary stress for the animal, and can potentially alter the hormones being measured. Because hormone metabolism is highly species‐specific, each assay must be validated for use in the species of interest. Validation of noninvasive steroid hormone assays has traditionally required the administration of relatively high doses of radiolabelled compounds (100 µCi or more of ¹⁴C labeled hormone) to permit subsequent detection of the excreted metabolites in the urine and feces. Accelerator mass spectrometry (AMS) is sensitive to extremely low levels of rare isotopes such as ¹⁴C, and provides a way to validate hormone assays using much lower levels of radioactivity than those traditionally employed. A captive Asian bull elephant was given 1 µCi of ¹⁴C‐testosterone intravenously, and an opportunistic urine sample was collected 2 hr after the injection. The sample was separated by HPLC and the ¹⁴C in the fractions was detected by AMS to characterize the metabolites present in the urine. A previously established HPLC protocol was used, which permitted the identification of fractions into which testosterone sulfate, testosterone glucuronide, and the parent compound testosterone elute. Results from this study indicate that the majority of testosterone excreted in the urine of the Asian bull elephant is in the form of testosterone sulfate. A small amount of testosterone glucuronide is also excreted, but there is no parent compound present in the urine at all. These results underscore the need for enzymatic hydrolysis to prepare urine samples for hormone assay measurement. Furthermore, they highlight the importance of proper hormone assay validation in order to ensure accurate measurement of the desired hormone. Although this study demonstrated the utility of AMS for safer validation of noninvasive hormone assays in nondomestic species, this methodology could also be applied to studies of nutrient metabolism and drug pharmakokinetics, both areas in great need of further study in wildlife species. Zoo Biol 29:760–766, 2010. © 2010 Wiley‐Liss, Inc.     

The savanna ecology of Kidepo Valley National Park, Uganda

Censuses indicated similar elephant densities in some habitats to those in certain other East African Parks where woodland has been almost completely destroyed. Giraffe densities were low. Herd structure indicated a healthy elephant population; but poor recruitment in the giraffe population. Policy for the national parks is discussed. It is suggested that it is not in the interests of Uganda or science to allow Kidepo Valley Park to become dominated by elephant in the same way as Kabalega and Rwenzori Park. Evidence that this is happening under current elephant use is strong. Cropping of two thirds of the elephant in the Narus valley is recommended. The revenue from cropping should be used for essential fire control measures and for a monitoring programme to ascertain optimum elephant numbers. Elephant should not be cropped in the absence of fire control.