Statural growth in known-age African elephants (Loxodonta africana)

The shoulder heights of 224 females and 170 males, and hindfoot length of 236 female and 217 male known-age African elephants ( Loxodonta africana ) were measured, and growth curves constructed for each measure of size. A linear relationship between foot length and shoulder height was confirmed in simultaneous measures of 97 males and 110 females. Growth curves demonstrated the typical sexual dimorphism in both foot length and shoulder height, with males growing more rapidly than females from birth onwards. The size dimorphism in foot length and shoulder height becomes marked by the age of 10 years, with males on average being 60–70 cm taller than females at 65 years. This size dimorphism is produced through faster growth which continues for longer than does that of females. The variance in growth rates is slightly greater for females than for males. It is proposed that female growth after puberty is affected by a trade-off between growth and reproduction, while males who deviate markedly from typical patterns of growth may be subject either to mortality or energetic constraints limiting their potential variance.     

Haematological studies on wild African elephants (Loxodonta africana)

Various haematological parameters have been measured in a sample of wild African elephants from two game parks in Uganda. The results are compared with those of other authors for the African and Indian elephant and other mammals. No intersex differences were found to occur but calves (1–5 years) differed from the population as a whole in several respects. Some values varied with season and this may be attributed to the effect of dehydration during the dry season. A number of elephants from one of the parks had abnormally high red cell numbers and the possibility is considered that this may reflect an incidence of habitat induced cardiovascular disease.     

Defaecation by African elephants (Loxodonta africana africana (Blumenbach))

A study of defaecation in the African elephant was carried out at the Voi headquarters of the Tsavo (East) National Park. Four orphaned animals aged between 1 and 10 years were observed for 4 days and 3 nights. During the day the time and weight of each individual's defaecation was recorded while at night only the time of defaecation. Details of all the records are presented. Analysis has shown that the amount of dung produced with each defaecation bears a similar characteristic to that of the growth curve of these animals. The rate of defaecation does not vary significantly with age. Records of defaecation arranged by time demonstrate an apparent periodicity with a low peak mid-morning and a high peak mid-afternoon. The potential use of this information in feeding and population studies is discussed.     

Effects of rainfall, host demography, and musth on strongyle fecal egg counts in African elephants (Loxodonta africana) in Namibia

Wild African elephants (Loxodonta africana) are commonly infected with intestinal strongyle parasites. Our objective was to determine baseline fecal strongyle egg counts for elephants in the northeast region of Etosha National Park, Namibia and determine if these numbers were affected by annual rainfall, elephant demography (age of individuals and composition of groups), and hormonal state of males. We found that matriarchal family group members have significantly higher fecal egg counts than male elephants (bulls). Among family group members, strongyle egg counts increased with age, whereas among bulls, strongyle egg counts decreased with age. Years of higher rainfall were correlated with decreased numbers of strongyle eggs among bulls. Finally, bulls were not affected by their physiologic (hormonal) status (musth vs. nonmusth). These results suggest that infection by strongyle parasites in Namibian African elephants is a dynamic process affected by intrinsic and extrinsic factors including host demography and rainfall.     

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.     

The sexually active states of free-ranging male African elephants (Loxodonta africana): defining musth and non-musth using endocrinology, physical signals, and behavior

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.     

African elephants (Loxodonta africana) recognize visual attention from face and body orientation

How do animals determine when others are able and disposed to receive their communicative signals? In particular, it is futile to make a silent gesture when the intended audience cannot see it. Some non-human primates use the head and body orientation of their audience to infer visual attentiveness when signalling, but whether species relying less on visual information use such cues when producing visual signals is unknown. Here, we test whether African elephants (Loxodonta africana) are sensitive to the visual perspective of a human experimenter. We examined whether the frequency of gestures of head and trunk, produced to request food, was influenced by indications of an experimenter''s visual attention. Elephants signalled significantly more towards the experimenter when her face was oriented towards them, except when her body faced away from them. These results suggest that elephants understand the importance of visual attention for effective communication.     

The relationship between social behaviour and habitat familiarity in African elephants (Loxodonta africana)

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.     

Estrous synchrony in a group of African elephants (Loxodonta africana) under human care

Synchrony of estrous, and consequently of conception and birth of young, may be of adaptive significance for certain mammals. Among the species in which estrous synchrony has been suspected several times are elephants, but clear evidence is still missing. We determined estrous cycles of African elephants (Loxodonta africana) (n=4) at the Vienna Zoo, Austria, between June 2003 and January 2006 by measuring serum progesterone levels from weekly blood samples. Except for the dominant female when she was intensively lactating, all animals showed clear cycles or progesterone release with a mean period of 105.3+/-15.37 days. For most of the study period, estrous cycles were asynchronous between females. However, after re-occurrence of the progesterone cycle in the dominant female following the first period of lactation, all four females showed high synchrony of progesterone release over the two subsequent cycles. Large changes in individual period lengths indicated that synchronization was due to the adjustment of cycle length in subdominants to that of the dominant female. We used a bootstrap procedure, based on resampling measured times of progesterone peaks, to determine if this apparent synchrony could have been caused by chance alone. This statistical analysis indicated that between-individual variances of the timing of progesterone peaks were much smaller that to be expected by chance (P=0.009). This finding represents the first evidence for estrous synchrony between elephants. We discuss various hypotheses to explain the biological function of cycle synchrony in elephants.     

Growth of the African elephant (Loxodonta africana)

The three-stage desk calculation of the von Bertalanffy equation to describe growth in height and weight with age in the elephant is compared with a new approach to calculating the three coefficients in the function by a computer. The two methods give different results with respect to the weight/age calculations. Theoretical von Bertalanffy equations calculated by both methods to describe growth in height and weight with age in the African elephant in Zambia are compared with previously published equations for the elephant in East Africa. Details are given of growth in height in two known-age African elephants, a female ‘Diksie’ and a male ‘Kartoum’. Theoretical growth in height curves for the female African and Asiatic elephant are compared. The coefficients K and / for growth in height are not transferable to the growth in weight equations. Inherent inaccuracies in the calculation of the coefficients in the von Bertalanffy equation are discussed, and it is concluded that in animals which have a long life-span such as the elephant, the equation serves as a purely empirical representation of weight-at-age data and that there is little biological significance in the parameters it contains. The computer-calculated curves give the best fit to the data. The regression of log age on log shoulder height from 2–20 years of age provides a more realistic approach to comparative growth studies. The increase in adrenal weight with age is linear. Tusk growth in relation to age and sex in Zambia is compared with East Africa. It is concluded that the tusks in Zambia are smaller and are more difficult to sex correctly than their East African counterparts, possibly a consequence of the Zambian elephant having a greater degree of tusk wear. Allometric growth is described with emphasis on the estimation of body weight from shoulder height. The most reliable estimates are obtained from a purely empirical representation of the data, a semilog plot of log body weight on shoulder height.     

Fecal 20-oxo-pregnane concentrations in free-ranging African elephants (Loxodonta africana) treated with porcine zona pellucida vaccine

Because of overpopulation of African elephants in South Africa and the consequent threat to biodiversity, the need for a method of population control has become evident. In this regard, the potential use of the porcine zona pellucida (pZP) vaccine as an effective means for population control is explored. While potential effects of pZP treatment on social behavior of African elephants have been investigated, no examination of the influence of pZP vaccination on the endocrine correlates in treated females has been undertaken. In this study, ovarian activity of free-ranging, pZP-treated African elephant females was monitored noninvasively for 1 yr at Thornybush Private Nature Reserve, South Africa, by measuring fecal 5α-pregnan-3β-ol-20-on concentrations via enzyme immunoassay. A total of 719 fecal samples from 19 individuals were collected over the study period, averaging 38 samples collected per individual (minimum, maximum: 16, 52). Simultaneously, behavioral observations were made to record the occurrence of estrous behavior for comparison. Each elephant under study showed 5α-pregnan-3β-ol-20-on concentrations rising above baseline at some period during the study indicating luteal activity. Average 5α-pregnan-3β-ol-20-on concentrations were 1.61 ± 0.46 μg/g (mean ± SD). Within sampled females, 42.9% exhibited estrous cycles within the range reported for captive African elephants, 14.3% had irregular cycles, and 42.9% did not appear to be cycling. Average estrous cycle duration was 14.72 ± 0.85 wk. Estrous behavior coincided with the onset of the luteal phase and a subsequent rise in 5α-pregnan-3β-ol-20-on concentrations. Average 5α-pregnan-3β-ol-20-on levels positively correlated with rainfall. No association between average individual 5α-pregnan-3β-ol-20-on concentrations or cyclicity status with age or parity were detected. Earlier determination of efficacy was established via fecal hormone analysis with no pregnancies determined 22 mo post-treatment and onward. Results indicate the presence of ovarian activity amongst pZP-treated female African elephants in 2 yr after initial immunization. Further study should now be aimed toward investigating the long-term effects of pZP vaccination on the reproductive function of female African elephants.     

Population dynamics of the African elephant (Loxodonta africana)

Recent studies on elephant populations from East Africa and from Zambia have suggested that as population density increases, so does the mean age at puberty and the mean calving interval. At the same time there is also an increase in the proportion of old females that are reproductively inactive. By constructing elephant population models, it is possible to investigate the extent to which these homeostatic mechanisms will regulate an elephant population. The models indicate that a change in the duration of the calving interval is more important as a population regulating mechanism than a change in the age at puberty, and that the proportion of old reproductively inactive females is of little significance. The importance of neonatal mortality in controlling population growth is emphasised by the models, and they also show that an annual population increase of 4% would be close to the maximum value.     

A population survey of forest elephants (Loxodonta africana cyclotis) in northern Congo

From February to April of 1989 transects were run in four forested areas of northern Congo (Brazzaville) to census elephant populations using dung counts. A total of 401.0 km of line transects was completed during the survey. An overall extrapolated density of 0.7 elephants/km² was obtained with a variation from 0.3 to 0.9 elephants/km². This study shows for the first time that elephants occur in high densities in a number of sites in northern Congo. When elephant dung density is plotted against distance from the nearest village for each transect a significant positive linear correlation results. Based on this correlation we hypothesize that the population of elephants in much of northern Congo is high. Local reports and our own observations indicate that elephant poaching is heavy in northern Congo and that the CITES ban on trade in African elephant ivory has had a limited effect on the level of poaching.     

Blood concentrations of gonadotrophins, prolactin and gonadal steroids in males and in non-pregnant and pregnant female African elephants (Loxodonta africana)

No seasonal variation in any of the hormones measured was apparent in males or females. Testosterone levels in males increased around puberty (10-11 years) and remained significantly higher in adult than prepubertal males. This was not accompanied by any significant change in levels of LH, FSH or prolactin. In non-pregnant females there was no apparent difference in levels of LH, FSH or prolactin with age. There was a significant increase in progesterone around puberty (12 years) but there was considerable overlap in values between prepubertal and adult females. During pregnancy, progesterone levels were significantly higher than in non-pregnant females with maximum levels occurring at mid-pregnancy (9-12 months). However, there was considerable overlap in values between non-pregnancy and pregnancy. Concentrations of LH and FSH decreased significantly during mid-pregnancy while prolactin levels increased dramatically during pregnancy; after 7 months of gestation until term levels were always at least 8 ng/ml greater than in any non-pregnant female. It is suggested that this consistent increase in plasma/serum levels of prolactin can be used to diagnose pregnancy in the elephant.