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.     

The movement of African elephants in a human-dominated land-use mosaic

Land outside of gazetted protected areas is increasingly seen as important to the future of elephant persistence in Africa. However, other than inferential studies on crop raiding, very little is understood about how elephants Loxodonta africana use and are affected by human-occupied landscapes. This is largely a result of restrictions in technology, which made detailed assessments of elephant movement outside of protected areas challenging. Recent advances in radio telemetry have changed this, enabling researchers to establish over a 24-h period where tagged animals spend their time. We assessed the movement of 13 elephants outside of gazetted protected areas across a range of land-use types on the Laikipia plateau in north-central Kenya. The elephants monitored spent more time at night than during the day in areas under land use that presented a risk of mortality associated with human occupants. The opposite pattern was found on large-scale ranches where elephants were tolerated. Furthermore, speed of movement was found to be higher where elephants were at risk. These results demonstrate that elephants facultatively alter their behaviour to avoid risk in human-dominated landscapes. This helps them to maintain connectivity between habitat refugia in fragmented land-use mosaics, possibly alleviating some of the potential negative impacts of fragmentation. At the same time, however, it allows elephants to penetrate smallholder farmland to raid crops. The greater the amount of smallholder land within an elephant's range, the more it was utilized, with consequent implications for conflict. These findings underscore the importance of (1) land-use planning to maintain refugia; (2) incentives to prevent further habitat fragmentation; (3) the testing and application of conflict mitigation measures where fragmentation has already taken place.     

圈养雌性非洲象妊娠期尿中性激素水平的变化

非洲象(Loxodonta africana)属于哺乳纲、长鼻目、象科、非洲象属,主要分布于非洲东部、中部、西部、西南部和东南部等广大地区,目前在世界大部分地区都有人工圈养.非洲象没有固定的发情季节,常年均可交配繁殖,雌兽的怀孕期为21 ～23个月,略长于亚洲象,是哺乳动物中孕期最长的动物.     

The impact of elephants on the marula tree Sclerocarya birrea

This study determined the abundance, density and population structure of the marula tree, Sclerocarya birrea, in three game reserves in South Africa, and assessed patterns and amounts of new and cumulative impact of elephants. Elephant feeding was very patchy so several attributes of individual trees, sampled transects and communities that might influence elephant herbivory were investigated. The incidence and type of elephant impact (bark, branch or stem breakage) were significantly related to tree diameter, but not to fruiting nor proximity to roads. At the transect level, elephant impact was influenced by density of marula trees, but was not influenced by proximity to roads, nor proportion of marula trees bearing fruits in the vicinity. At the community level, elephant impact was higher on reserves with higher total marula densities. Fourfold differences in elephant densities (0.08–0.30 elephants km^?2) did not explain marula consumption: the percentage of trees with branch damage was similar across reserves and bark damage was inversely proportional to elephant density. Variation across reserves may reflect local and landscape‐level marula tree abundance, differences in alternative food plants and individual feeding habits. The recorded levels of impact appeared to be sustainable because mortality rates were low, affected trees often recovered, and small trees were not preferentially preyed upon.     

The use of low-frequency vocalizations in African elephant(Loxodonta africana) reproductive strategies

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.     

Morphology of the female reproductive organs of the African lion (Panthera leo)

The topography and splanchnology of the reproductive organs of the African lioness were studied and described in situ and after removal. The kidneys were located far caudally in relation to the thirteenth ribs. The suspensory ligament was very well developed, originated in a fan‐like manner from the dorsolateral abdominal wall lateral to the kidney and extended up to a few centimetres cranial to the kidney. The proper ligament of the ovary as well as the round ligament was well developed. The round ligament inserted on the medial femoral fascia. The left ovary was bigger than the right. The ovarian bursa had a short mesosalpinx that did not cover any part of the ovary and the fimbriae extended along the entire length. The urethral tuberculum as well as the urethral crest were well developed. The left uterine horn was longer than the right. The uterine tube was found to open directly into the tip of the uterine horn and not onto a papilla. The reproductive organs of the lioness resembled those of the domestic cat and dog but with some major differences.     

Physiology and endocrinology of zinc accumulation during the female squirrelfish reproductive cycle

Females of the squirrelfish family (Holocentridae) accumulate higher levels of hepatic zinc than any other studied animal. This accumulation is accompanied by high expression of the zinc-binding protein, metallothionein (MT), and is strongly correlated to the onset of sexual maturity. In an attempt to further characterize the timeframe of this accumulation, and to possibly discern any potential mediators, we examined the physiology and endocrinology of the yearly reproductive cycle of mature female squirrelfish. There are two separate reproductive events during the year in December-January and again in March-April, as evidenced by peaks in ovarian growth, VTG production, steroid levels, zinc accumulation and redistribution. Increased hepatic zinc seems to be preceded by a necessary increase in MT, but this was not clearly correlated to plasma 17beta-estradiol, testosterone, or progesterone levels. The plasma zinc protein vitellogenin (VTG) is one, but probably not the predominant, vehicle for the transport of hepatic zinc to the ovary.     

A Meta-Analysis of the Impact of African Elephants on Savanna Vegetation

Abstract: Large herbivores such as elephants (Loxodonta africana) apparently have a negative impact on woody vegetation at moderate to high population densities. The confounding effects that fire, drought, and management history have may complicate assignment of such impacts to herbivory. We reviewed 238 studies published over 45 years and conducted a meta-analysis based on 21 studies that provided sufficient information on response of woody vegetation to elephants. We considered size and duration of studies, elephant densities, rainfall, fences, and study outcomes in our analysis. We detected a disproportionate citation of 20 published studies in our database, 15 of which concluded that woody vegetation responded negatively to elephants. Our analysis showed that high elephant densities had a negative effect on woody vegetation but that rainfall and presence of fences influenced these effects. In arid savannas, woody vegetation always responded negatively to elephants. In transitional savannas, an increase in elephant densities did not influence woody vegetation response. In mesic savannas, negative responses of woody vegetation increased when elephants occurred at higher densities, whereas elephants confined by fences also had more negative effects on woody plants than elephants that were not confined. Our analysis suggested that rainfall and fences influenced elephant density related impact and that research results were often site-specific. Local environmental conditions and site-specific objectives should be considered when developing management actions to curb elephant impacts on woody vegetation.     

西双版纳三岔河地区野生亚洲象的个体识别、种群数量和活动特点

2004 年10 月至2005 年10 月,在云南西双版纳三岔河野象谷用直接观察、拍照、摄像等方法直接记录野生亚洲象的形态学特征,通过耳、门齿、背、尾、疤痕、面部骨骼等特征对经过本地区的象群和独象进行个体识别并建立个体信息库。直接观测记录野象524 头次,通过照片资料和访问间接记录289 头次,识别为46 头个体,由7 个家族(庭)群共38 头和8 头单独活动的雄象组成;发现象群和独象的活动具有季节性;不同象群在野象谷的活动频次差异显著;象群大小和其在本地区的活动频率之间呈明显的正相关。     

The environmental context for the origins of modern human diversity: a synthesis of regional variability in African climate 150,000-30,000 years ago

We synthesize African paleoclimate from 150 to 30 ka (thousand years ago) using 85 diverse datasets at a regional scale, testing for coherence with North Atlantic glacial/interglacial phases and northern and southern hemisphere insolation cycles. Two major determinants of circum-African climate variability over this time period are supported by principal components analysis: North Atlantic sea surface temperature (SST) variations and local insolation maxima. North Atlantic SSTs correlated with the variability found in most circum-African SST records, whereas the variability of the majority of terrestrial temperature and precipitation records is explained by local insolation maxima, particularly at times when solar radiation was intense and highly variable (e.g., 150-75 ka). We demonstrate that climates varied with latitude, such that periods of relatively increased aridity or humidity were asynchronous across the northern, eastern, tropical and southern portions of Africa. Comparisons of the archaeological, fossil, or genetic records with generalized patterns of environmental change based solely on northern hemisphere glacial/interglacial cycles are therefore imprecise.We compare our refined climatic framework to a database of 64 radiometrically-dated paleoanthropological sites to test hypotheses of demographic response to climatic change among African hominin populations during the 150-30 ka interval. We argue that at a continental scale, population and climate changes were asynchronous and likely occurred under different regimes of climate forcing, creating alternating opportunities for migration into adjacent regions. Our results suggest little relation between large scale demographic and climate change in southern Africa during this time span, but strongly support the hypothesis of hominin occupation of the Sahara during discrete humid intervals ∼135-115 ka and 105-75 ka. Hominin populations in equatorial and eastern Africa may have been buffered from the extremes of climate change by locally steep altitudinal and rainfall gradients and the complex and variable effects of increased aridity on human habitat suitability in the tropics. Our data are consistent with hominin migrations out of Africa through varying exit points from ∼140-80 ka.     

Influencing factors on the foot health of captive Asian elephants (Elephas maximus) in European zoos

Pathological lesions of feet occur frequently in captive elephant populations. To improve foot health, it is important to identify risk factors associated with such pathologies. Several previous studies have analyzed potentially influencing factors but were limited, for example, by small sample sizes. This study analyzed the relationship between 87 independent variables and the foot health score of 204 Asian elephants (Elephas maximus) in European zoos using bivariate correlation, multivariable regression models, and principal component analysis (PCA). Correlation and regression tests revealed significant results for 30 different variables, mainly with small effect sizes. Only three variables were significant in more than one test: sex, time spent indoors, and time spent on hard ground, with lower scores (i.e. less or less severe pathological lesions) in females, and when less time is spent indoors or on hard ground. Due to small effect sizes and differing results of the statistical tests, it is difficult to determine which risk factors are most important. Instead, a holistic consideration appears more appropriate. A biplot of the PCA shows that factors representing more advanced husbandry conditions (e.g. large areas, high proportions of sand flooring) were associated with each other and with decreased foot scores, whereas indicators of more limited conditions (e.g. high proportions of hard ground, much time spent indoors) were also associated with each other but increased the foot score. In conclusion, instead of resulting from just one or two factors, reduced foot health might be an indicator of a generally poorer husbandry system.     

社群压力对圈养雌性亚洲象动情的影响

亚洲象是中国一级濒危保护动物,改善圈养亚洲象的繁殖状况是亚洲象保护工作的重要内容之一。本研究旨在通过分析社群压力对圈养雌性亚洲象应激和动情状况的影响,从而探究圈养雌性亚洲象长期不动情的原因。通过放射性免疫法检测雌性亚洲象尿液中皮质醇、孕酮和雌二醇水平以及粪便中皮质醇和雌二醇水平,以雌激素水平反映亚洲象的动情状况,以皮质醇水平反映亚洲象的应激状态,以争斗行为和友好行为的发生情况反映圈养群体的社群压力。研究发现：(1)处于优势地位的个体表现出更多的斗争行为,处于从属地位的个体受到更多的攻击行为;(2)优势个体的皮质醇水平显著高于处于从属地位个体的皮质醇水平;(3)优势个体的雌二醇和孕酮具有小幅周期性波动,而从属个体的雌二醇和孕酮水平没有表现出明显的波动。从本研究结果可以看出圈养亚洲象群体中优势个体对从属个体的行为抑制可能是导致从属个体雌激素水平低,长期不能参与繁殖的原因。     

Effects of reproductive and social context on vocal communication in captive female African elephants (Loxodonta africana)

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. Zoo Biol 0:1–17, 2005. © 2005 Wiley‐Liss, Inc.     

Investigation of individual and group variability in estrous cycle characteristics in female Asian elephants (Elephas maximus) at the Oregon Zoo

Evaluating ovarian cycle activity through longitudinal progestagen monitoring is important for optimizing breeding management of captive elephants and understanding impact of life events (births, deaths, and transfers) on reproductive function. This study summarized serum progestagen profiles for eight Asian mainland elephants (Elephas maximus indicus) and one Bornean elephant (E. maximus borneensis) at the Oregon Zoo over a 20-yr interval, and represents the longest longitudinal dataset evaluated to date. Estrous cycle characteristics were more varied than previously reported for this species, with an overall duration of 12 to 19 wk, luteal phase duration of 4 to 15 wk, and follicular phase duration of 2 to 12 wk. In general, there was more cycle variability across than within individual elephants. Compared with other elephants in the group, the Borneo female exhibited consistently longer cycle lengths, higher progestagen concentrations, and greater cycle variability; however, it is not known if this represents a subspecies or an individual difference. Cycle durations did not appear to change over time or with age, and the first pubertal cycle was similar to subsequent cycles. Variability in duration of the follicular phase was greater than that of the luteal phase. In addition, there was a significant negative relationship between luteal and follicular phase durations, suggesting a possible regulatory role of the follicular phase in maintaining a relatively consistent cycle duration within individuals. Overall, we found these elephants to be highly resilient in that major life events (births, deaths, and changes in herd structure) had minimal effect on cycle dynamics over time. In conclusion, the higher range in cycle phase characteristics is likely because of the larger number of elephants studied and longer duration of longitudinal monitoring, and may be more representative of the captive population as a whole. Furthermore, identification of significant interanimal variability suggests that understanding the complexities of herd reproductive characteristics could facilitate development of more effective institution-specific breeding management strategies.     

Ultrasonographic and laparoscopic evaluation of the reproductive tract of the captive female African lion (Panthera leo)

The use of transabdominal ultrasonography to assess the oestrous cycle has not been previously described in the African lion (Panthera leo). Twelve sexually mature lionesses and five female cubs had their reproductive organs assessed by transabdominal ultrasound. Ovarian findings were compared to laparoscopic findings while performing laparoscopic ovariectomy or salpingectomy. Vaginal cytology was performed and serum progesterone levels were determined. By combining all data the oestrous cycle stage of each lion was determined. One lion was far pregnant and was not operated on. In adults a uterine body could be seen ultrasonographically in 67% of lions while mural structures could be distinguished in 44% of lions. Five uterine horns could be seen in 3 lions. In 12 adults 10 ovaries were found of which eight had discernable follicles or luteal structures. During laparoscopy 12 active ovaries were seen with luteal structures seen in 11 ovaries and follicles in 2 ovaries. Using laparoscopy as the gold standard, ultrasonography had a sensitivity of 66% and specificity of 83% to detect ovarian reproductive activity. Two uterine cysts and a cluster of periovarian cysts were seen in three different lions. Three lions were pregnant, two were in oestrus, three in a luteal phase (dioestrus), and four were in anoestrus. Transabdominal ultrasound in combination with serum progesterone levels and vaginal cytology can be used to assess ovarian cyclical activity with reasonable accuracy in captive bred lions.     

Contemporary and historical impacts of megaherbivores on the population structure of tree euphorbias in South African subtropical thicket

African elephant and black rhinoceros – both megaherbivores – impact negatively on the abundance of succulent plants, including tree succulents, in South Africa's subtropical thicket. We sampled 35 sites in subtropical thicket to assess historical and contemporary impacts of African elephant and black rhinoceros on the population structure of three species of succulent tree euphorbia. Population age structures were highly variable and showed no species-specific effects. Almost half the sites had growing populations dominated by young individuals. Sites having contemporary impacts had significantly fewer individuals in the 30–75 years age range, but this effect declined with increasing terrain slope. Eighty-one percent of sites assumed to have been impacted by megaherbivores historically had individuals that predated rhino and elephant extirpation in the region. This and other population age structure data indicate that megaherbivores and tree euphorbias coexisted historically on terrain of low relief. However, in areas not subject to contemporary impacts, euphorbia populations are likely to be much higher now than historically, owing to population relaxation after the extirpation of megaherbivores in the mid 1800s. Given the sensitivity of tree euphorbias to megaherbivore impacts, managers should consider monitoring the population structure of these species in order to identify thresholds of potential concern regarding megaherbivore impacts.     

Patterns of elephant impact on woody plants in the Hluhluwe-Imfolozi park, Kwazulu-Natal, South Africa

This study identifies patterns of elephant Loxodonta africana africana impacts upon tree species and woody plant communities in Hluhluwe-Imfolozi Park, a South African savannahs/woodlands area. Elephants were reintroduced there from 1981, following more than 80 years of absence. Data were collected in 2003 on elephant impact on woodland in the Park. Different vegetation types were susceptible to different types and levels of damage by elephants, suggesting that elephants will not homogenize the vegetation. Elephants targeted larger stems for all types of damage, with a strong preference for some of the less abundant species such as Albizia versicolor (breaking and toppling) and Cordia caffra and Schotia brachypetala (debarking). Elephant impacts tended to be distributed evenly across the park landscape, irrespective of stem density or proximity to permanent water. Overall, elephants have little impact on slowing or reversing the spread of undesirable woody species, but are having a marked impact on certain less common tree species and larger tree size-classes in the Hluhluwe-Imfolozi Park.