Social and environmental impacts on sleep in captive Asian elephants (Elephas maximus)

Modern zoos strive to improve standards of animal management, husbandry and welfare of their animals as part of a continual evaluation process. Elephants (Elephantidae) have received particular attention in recent years due to the challenge of providing environments which promote natural behavior and opportunities for social interaction. A number of measures have been proposed to measure wellbeing, with sleep quality increasingly being used. Sleep is a vital aspect of life for cell replenishment as well as optimal development of young. Sleep deprivation can lead to immunosuppression and illness; therefore animal managers have a responsibility to ensure they reduce the potential for disturbance through noise, light, or other environmental factors. The social environment also plays an essential role in wellbeing, particularly for species that live in multi-generational family units. In this study the nocturnal behavior of a multi-generational captive herd was observed to determine impacts of husbandry changes on sleep duration and bout length (measured as recumbent rest). As expected, average total duration of sleep was higher in younger elephants and rates were comparable to those reported in other studies of Asian elephants. Overnight access to an outdoor paddock in warmer weather increased overall average bout length of sleep in the herd. Average total duration of sleep also increased for the herd following the movement of an unrelated adult female who had previously shown weak bonds with other herd members. This indicates that social compatibility is a vital component of elephant welfare, impacting not only behavioral interactions but sleep quality and duration.     

The behavioral effects of exhibit size versus complexity in African elephants: A potential solution for smaller spaces

Population‐level analyses suggest that habitat complexity, but not necessarily space availability, has important welfare outcomes for elephants in human care. At the Dallas Zoo, the opening of a new exhibit complex allowed us to measure the behavior of two female African elephants across three treatments to evaluate the independent effects of complexity and space. Preoccupancy observations were conducted in the elephants’ older exhibit, which consisted of a smaller, more simple yard (630 m²). Subsequent postoccupancy observations measured behavior in two different spaces in the new exhibit: a larger, complex yard (15,000 m²), and a smaller, but complex yard (1,520 m²). The elephants’ overall activity levels were greater in complex habitats, regardless of their size. Similar effects of habitat complexity oversize were observed with greater rates of foraging and lower rates of being stationary. Furthermore, elephants were out of view of visitors significantly more in the small, simple yard compared to either of the more complex habitats. However, exhibit size affected the incidence of stereotypic behavior (with lower rates of stereotypy in the larger exhibit compared to the smaller yards) and investigatory behavior (elephants investigated their environments more with increasing size and complexity). Behavioral diversity also increased with exhibit size and complexity. These results indicate that space availability alone is not sufficient to enhance the behavioral welfare of zoo elephants. Therefore, facilities with limited space can still encourage species‐appropriate behaviors and improved welfare for the elephants in their care by converting a small, simple area into a more complex habitat.     

The Social and Ecological Integration of Captive-Raised Adolescent Male African Elephants (Loxodonta africana) into a Wild Population

Background

A rapid rise in the number of captive African elephants (Loxodonta africana) used in the tourism industry in southern Africa and orphaned elephants in human care has led to concerns about their long-term management, particularly males. One solution is to release them into the wild at adolescence, when young males naturally leave their herd. However, this raises significant welfare concerns: little is known about how well released elephants integrate into wild populations and whether they pose a greater threat to humans than wild elephants. We document the release of three captive-raised adolescent male African elephants in the Okavango Delta, Botswana.

Methodology/Principal Findings

Despite having been part of a herd of working elephants for at least eight years, the three males progressively integrated into the complex fission-fusion society of wild bull elephants. In the three years following release, they showed no tendency to be closer to human habitation, and there were no significant differences between wild and captive-raised adolescent males in the total number of social interactions, size of ranges and habitat use. However, the captive-raised elephants sparred less and vocalised more, and spent more time alone and in smaller social groups. Thereafter the released elephants continued to expand their ranges and interact with both mixed-sex herds and males. One male was shot by farmers 94 months after release, along with ten wild elephants, on a ranch outside the protected area.

Conclusions/Significance

We show that captive-raised adolescent male elephants can integrate into a wild population. Long-term studies are required to determine the longevity, breeding success, and eventual fate of released male elephants, but we identified no significant short-term welfare problems for the released elephants or recipient population. Release of captive-raised mammals with complex social systems is a husbandry option that should be explored further.     

Spatial distribution and movements of elephants (Loxodonta africana) in the northern Namib Desert region of the Kaokoveld, South West Africa/Namibia

The home ranges, space-use patterns, seasonal distribution, long-distance movements, water-related movements and movement patterns of elephants ( Loxodonta africana ) residing in the northern Namib Desert region of the Kaokoveld, South West Africa/Namibia, are discussed. All movements of the desert-dwelling elephants, including occasional wanderings up to 195 km, were confined to the northern Namib Desert and no evidence could be found of any movements including migrations to and from the Etosha National Park. Home range size varied from 1763 to 2944 km² and the main feature of the spatial distribution of these elephants is the marked fidelity to individual home ranges. Seasonal distribution and space-use patterns within individual home ranges coincide with seasonal climatic changes and the corresponding changes in food and water availability. The elephant's normal movements and migrations were confined to individual home ranges and no mass population migrations took place. Their mobility and their ability to go up to four days without drinking water enabled them to utilize food sources up to 70 km away from waterholes and, together with their intimate knowledge about resource distribution within their home ranges, are regarded as key factors in their survival in the desert. The elephants' spatial distribution and movement patterns indicate a natural population segregation from other elephant populations in the Kaokoveld, clearly demarcating the desert-dwelling elephants as a distinct population with their own movement patterns and geographic range.     

Elucidating the significance of spatial memory on movement decisions by African savannah elephants using state–space models

Spatial memory facilitates resource acquisition where resources are patchy, but how it influences movement behaviour of wide-ranging species remains to be resolved. We examined African elephant spatial memory reflected in movement decisions regarding access to perennial waterholes. State–space models of movement data revealed a rapid, highly directional movement behaviour almost exclusively associated with visiting perennial water. Behavioural change point (BCP) analyses demonstrated that these goal-oriented movements were initiated on average 4.59 km, and up to 49.97 km, from the visited waterhole, with the closest waterhole accessed 90% of the time. Distances of decision points increased when switching to different waterholes, during the dry season, or for female groups relative to males, while selection of the closest waterhole decreased when switching. Overall, our analyses indicated detailed spatial knowledge over large scales, enabling elephants to minimize travel distance through highly directional movement when accessing water. We discuss the likely cognitive and socioecological mechanisms driving these spatially precise movements that are most consistent with our findings. By applying modern analytic techniques to high-resolution movement data, this study illustrates emerging approaches for studying how cognition structures animal movement behaviour in different ecological and social contexts.     

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.     

Interrelationship of serum testosterone, dominance and ovarian cyclicity status in female African elephants

The African elephant population in North American zoos is not self-sustaining, in part due to the prevalence of ovarian acyclicity. While little is known about the cause of this condition, earlier research has shown that females without cyclic corpus luteum (CL) function rank higher in the dominance hierarchy than females with cyclic CL function. The goal of this study was to measure longitudinal serum testosterone concentrations in captive female African elephants to determine if there is a relationship among serum testosterone concentrations, social dominance rank and ovarian cyclicity status. Weekly blood samples from 49 female African elephants (24 having and 25 not having cyclic CL function at 22 facilities) were collected over a 12-month period and analyzed for serum testosterone using an enzymeimmunoassay. A progesterone radioimmunoassay was used to quantify serum progestagen concentrations and categorize ovarian cyclicity status. The dominance hierarchy of individual elephants within each herd was assessed by a written temperament survey, which identified 19 dominant, 15 middle and 15 subordinate females. No clear patterns of serum testosterone secretion were observed in females with and without cyclic CL function. Furthermore, no significant relationships were found among serum testosterone concentrations, dominance rank, and ovarian cyclicity status. These data suggest that increased circulating testosterone concentrations are not associated with greater rates of ovarian acyclicity or dominance status in captive female African elephants.     

Response of African Elephants (Loxodonta africana) to Seasonal Changes in Rainfall

The factors that trigger sudden, seasonal movements of elephants are uncertain. We hypothesized that savannah elephant movements at the end of the dry season may be a response to their detection of distant thunderstorms. Nine elephants carrying Global Positioning System (GPS) receivers were tracked over seven years in the extremely dry and rugged region of northwestern Namibia. The transition date from dry to wet season conditions was determined annually from surface- and satellite-derived rainfall. The distance, location, and timing of rain events relative to the elephants were determined using the Tropical Rainfall Measurement Mission (TRMM) satellite precipitation observations. Behavioral Change Point Analysis (BCPA) was applied to four of these seven years demonstrating a response in movement of these elephants to intra- and inter-seasonal occurrences of rainfall. Statistically significant changes in movement were found prior to or near the time of onset of the wet season and before the occurrence of wet episodes within the dry season, although the characteristics of the movement changes are not consistent between elephants and years. Elephants in overlapping ranges, but following separate tracks, exhibited statistically valid non-random near-simultaneous changes in movements when rainfall was occurring more than 100 km from their location. While the environmental trigger that causes these excursions remains uncertain, rain-system generated infrasound, which can travel such distances and be detected by elephants, is a possible trigger for such changes in movement.     

云南思茅亚洲象对栖息地的选择与利用

以思茅市境内栖居的一个由5 头雌象(3 头成年体, 1 头亚成体, 1 头幼体) 组成的象群为研究对象, 利用痕迹追踪、样线调查和村庄调查等方法对野生亚洲象的栖息地选择和行为进行了初步研究。在栖息地中共记录到野生高等植物457 种, 组成5 种植被类型。旱季象群的活动区域面积是35.67 km² , 该活动区域中有3 个核心活动区, 面积分别是3.65 km² , 2.79 km² 和3.29 km² , 象群对此3 个活动核心区域循环利用; 在旱季亚洲象经常取食19种野生植物, 其中主要取食的有8 种, 还取食7 种农作物, 对农作物有一定依赖性, 这种依赖的程度随着农田周围森林中野生食物丰富程度的增加而降低; 在季雨林中亚洲象主要取食的植物种类最丰富。雨季象群的活动区域面积是18.42 km² , 该区域中只有1 个核心活动区, 面积9.08 km² 。雨季亚洲象只取食5 种野生植物, 另取食5 种农作物, 农作物为其雨季主要食物。研究结果表明, 思茅地区亚洲象栖息地中的野生食物可能不足, 人类活动对亚洲象的生存干扰较为严重, 亚洲象的行为表现出对栖息地内食物条件和人类干扰的适应。建议尽快遏止日益严重的栖息地片断化趋势, 恢复自然植被并为亚洲象提供充足的野生食物是保护该物种的关键。     

Vigilance behavior of Pyrenean chamois Rupicapra pyrenaica pyrenaica: Effect of sex and position in the herd

The Pyrenean chamois Rupicapra pyrenaica pyrenaica is a mountain-dwelling ungulate with an extensive presence in open areas. Optimal group size results from the trade off between advantages (a reduction in the risk of predation) and disadvantages (competition between members of the herd) of group living. In addition, advantages and disadvantages of group living may vary depending on the position of each individual within the herd. Our objective was to study the effect of central vs. Peripheral position in the herd on feeding and vigilance behavior in male and female Pyrenean chamois and to ascertain if a group size effect existed. We used focal animal sampling and recorded social interactions when a focal animal was involved. With males, vigilance rate was higher in the central part of the group than at the periphery, probably due to a higher density of animals in the central part of the herd and a higher probability of being disturbed by conspecifics. With females, vigilance rate did not differ according to position in the herd. Females spent more time feeding than males, and males showed a higher frequency of the vigilance behavior than females. We did not observe a clear relationship between group size and vigilance behavior. The differences in vigilance behavior might be due to social interactions.     

Movements,reproductive seasonality,and fisheries interactions in the whitetip reef shark (<Emphasis Type="Italic">Triaenodon obesus</Emphasis>) from community-contributed photographs

Despite being a common apex-level predator on coral reefs throughout the tropical Indo-Pacific, surprisingly little is known about whitetip reef shark (Triaenodon obesus) movements and biology. This study used photo-identification from community-contributed photographs to reveal patterns in movements, reproductive biology, and fisheries interactions in this species that have not been previously revealed through more traditional methods. At least 178 individual sharks were identified, and 26 movements were observed. These included movement distances of up to 26.4 km, movement rates of up to 3.27 km/day (9.8 km in 3 days), and movements that required the transit of a 140 m deep channel. Other animals showed high philopatry, being re-sighted at the same locality on multiple occasions (up to 13 sightings for one individual) over periods of up to 7 years. Females showed higher philopatry than males and were more likely than males to be found at shallow (<10 m depth) localities throughout the year. The proportion of male sightings at shallow localities was significantly higher in April and May than other months of the year, possibly due to males coming into the shallows to mate with females. A peak in sightings of late-term females followed by an abrupt decline suggests that pupping season is May into early June, and two females were observed pregnant in consecutive years despite evidence that the gestation period is approximately 1 year for this species. Nine percent of animals carried fishing tackle or exhibited jaw injuries associated with fishery interactions, with multiple individuals found dead after being hooked by fishers.     

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.     

What if it gets crowded? Density‐dependent tortuosity in individual movements of a Neotropical mammal

Effects of density dependence on animal movements have received much attention in ecology, but it is still debated to what extent dispersal and movements in general are density dependent, and their potential contribution to population regulation processes. Here, we determine the occurrence and nature of density dependence in the movements of a Neotropical marsupial, the black‐eared opossum Didelphis aurita Wied‐Neuwied 1826. Using spool‐and‐line tracking devices, we estimated the tortuosity of fine‐scale movements of 149 individuals by their fractal dimension D. We evaluated the relative importance of population size, reproductive or climatic seasons and reproductive maturity of individuals as determinants of movement tortuosity, using a model selection approach. Population size was the most important determinant of movement tortuosity, with season (climatic seasons for females, reproductive seasons for males) and reproductive maturity as secondary but also important variables. We detected a positive density‐dependent effect on movement tortuosity, resulting in more intensive use of areas by individuals during periods of high population size. This positive association between movement tortuosity and population size is more likely to result from intraspecific competition, which forces individuals to explore their environment more intensively during high‐density periods. Therefore, despite being density dependent, movements in D. aurita apparently do not contribute to population regulation mechanisms.     

Roadless wilderness area determines forest elephant movements in the Congo Basin

A dramatic expansion of road building is underway in the Congo Basin fuelled by private enterprise, international aid, and government aspirations. Among the great wilderness areas on earth, the Congo Basin is outstanding for its high biodiversity, particularly mobile megafauna including forest elephants (Loxodonta africana cyclotis). The abundance of many mammal species in the Basin increases with distance from roads due to hunting pressure, but the impacts of road proliferation on the movements of individuals are unknown. We investigated the ranging behaviour of forest elephants in relation to roads and roadless wilderness by fitting GPS telemetry collars onto a sample of 28 forest elephants living in six priority conservation areas. We show that the size of roadless wilderness is a strong determinant of home range size in this species. Though our study sites included the largest wilderness areas in central African forests, none of 4 home range metrics we calculated, including core area, tended toward an asymptote with increasing wilderness size, suggesting that uninhibited ranging in forest elephants no longer exists. Furthermore we show that roads outside protected areas which are not protected from hunting are a formidable barrier to movement while roads inside protected areas are not. Only 1 elephant from our sample crossed an unprotected road. During crossings her mean speed increased 14-fold compared to normal movements. Forest elephants are increasingly confined and constrained by roads across the Congo Basin which is reducing effective habitat availability and isolating populations, significantly threatening long term conservation efforts. If the current road development trajectory continues, forest wildernesses and the forest elephants they contain will collapse.     

Estimating the defaecation rate of the African forest elephant (Loxodonta cyclotis) in Banyang-Mbo Wildlife Sanctuary, south-western Cameroon

Defaecation rate is an important parameter in the conversion of dung counts to elephant numbers, yet estimates of elephant defaecation rates are available for a few sites, especially rainforest sites. The Banyang‐Mbo Wildlife Sanctuary in south‐western Cameroon is no exception in this regard. We tracked a radio‐collared herd of four forest elephants, counted their fresh droppings in two 24‐h periods each month in 36 consecutive months. Our results show a mean defaecation rate of 15.9 [95% CI (12.2–19.7)] dung piles per elephant per day. We found significant monthly and inter‐annual variation in defaecation rates with implications for their predictability and forest elephant surveys. In particular, our results show that mean defaecation rates are higher for the wettest period of the year than the relatively dry months.     

Water consumption and watering behavior of Przewalski horses (Equus ferus przewalskii) in a semireserve

Water consumption by 12 female Przewalski horses was automatically measured in a semireserve for 17 months. Average daily consumption on a yearly basis was between 2.4 and 8.3 liters, but significant individual variation in water consumption and drinking frequency was recorded throughout the observation period. Individual differences were more clearly detectable during periods of hot and dry weather. Individuals with high water demands initiated herd movements toward watering places independent of their social rank. Discussed as possible reasons for individual differences in water demand are: acquired preferences, distinct food selection, and genetically determined physiological differences. Watering behavior should not be regarded as an important factor only in re-introduction projects, but also with respect to animal welfare in general. Zoo Biol 17:181–192, 1998. © 1998 Wiley-Liss, Inc.     

A Case Study: Fecal Corticosteroid and Behavior as Indicators of Welfare During Relocation of an Asian Elephant

This study was a preliminary investigation of an enzyme immunoassay for measuring fecal glucocorticoid metabolites in a male Asian elephant (Elephas maximus) by investigating changes in behavior and cortisol metabolite excretion associated with a putative stressful event. The study collected fecal samples for 10 days prior to, and 10 days after, 24-hr transport and relocation of the elephant to a new herd. The study measured cortisol metabolites using 2 enzyme immunoassays indicating a 389% and 340% increase in cortisol metabolite excretion following relocation. Maximal cortisol metabolite excretion occurred 2 days after relocation and remained elevated during establishment of the new herd. Stereotypic behavior increased approximately 400% after relocation. The relocation disturbed sleep patterns, the elephant spent less time sleeping during the night, and the elephant slept standing up. These results provide preliminary evidence that noninvasive monitoring of fecal cortisol metabolites can be used to investigate adrenal activity in Asian elephants and may be a safe, practical, and accurate welfare indicator.     

Density‐dependent space use affects interpretation of camera trap detection rates

Camera traps (CTs) are an increasingly popular tool for wildlife survey and monitoring. Estimating relative abundance in unmarked species is often done using detection rate as an index of relative abundance, which assumes that detection rate has a positive linear relationship with true abundance. This assumption may be violated if movement behavior varies with density, but the degree to which movement behavior is density‐dependent across taxa is unclear. The potential confounding of population‐level relative abundance indices by movement would depend on how regularly, and by what magnitude, movement rate and home‐range size vary with density. We conducted a systematic review and meta‐analysis to quantify relationships between movement rate, home‐range size, and density, across terrestrial mammalian taxa. We then simulated animal movements and CT sampling to test the effect of contrasting movement scenarios on CT detection rate indices. Overall, movement rate and home‐range size were negatively correlated with density and positively correlated with one another. The strength of the relationships varied significantly between taxa and populations. In simulations, detection rates were related to true abundance but underestimated change, particularly for slower moving species with small home ranges. In situations where animal space use changes markedly with density, we estimate that up to thirty percent of a true change in relative abundance may be missed due to the confounding effect of movement, making trend estimation more difficult. The common assumption that movement remains constant across densities is therefore violated across a wide range of mammal species. When studying unmarked species using CT detection rates, researchers and managers should explicitly consider that such indices of relative abundance reflect both density and movement. Practitioners interpreting changes in camera detection rates should be aware that observed differences may be biased low relative to true changes in abundance. Further information on animal movement, or methods that do not depend on assumptions of density‐independent movement, may be required to make robust inferences on population trends.     

How can leadership processes in European bison be used to improve the management of free-roaming herds

For several years, several wild species are experiencing a comeback in Europe, especially large herbivores. Despite the good news, the increased number of wild herbivores also potentially increases human-wildlife conflicts due to crop damages and road traffic accidents where such species have been involved in previously. In this context, studying the movements of animal groups is particularly relevant for managing their use of space and ensure sufficient cohabitation with human populations. We thus studied the European bison (Bison bonasus), a species concerned in several rewilding programs, to understand to what extent individual characteristics and social relationships are involved in the leadership process and how these factors influence the success of group movements. We observed a herd of 14 individuals in Zuid-Kennemerland National Park (the Netherlands) and analysed the initiation phase of spontaneous group movements. We showed that leadership was distributed throughout the group but the two oldest females showed higher propensity to initiate and to be followed. They seemed to have a high social power and a strong capacity to federate probably due to their dominance rank and their knowledge of the areas. The joining process was faster when animals were grouped before departure and when the initiator moved in the direction towards which the majority of individuals was oriented. Individuals joined the movement according to a selective mimetic process, based on kinship. These results, combined with virtual fence device, could be used to manage herd movements in a given territory and thus, might be an innovative tool to decrease human-wildlife conflicts.     

A Comparison of Walking Rates Between Wild and Zoo African Elephants

With increased scrutiny surrounding the welfare of elephants in zoological institutions, it is important to have empirical evidence on their current welfare status. If elephants are not receiving adequate exercise, it could lead to obesity, which can lead to many issues including acyclicity and potentially heart disease. The goal of the current study was to compare the walking rates of elephants in the wild versus elephants in zoos to determine if elephants are walking similar distances relative to their wild counterparts. Eleven wild elephants throughout different habitats and locations in Botswana were compared to 8 elephants at the San Diego Zoo Safari Park. Direct comparisons revealed no significant difference in average walking rates of zoo elephants when compared with wild elephants. These results suggest that elephants at the San Diego Zoo Safari Park walk similar rates to those of wild elephants and may be meeting their exercise needs.