Elephant survival,rainfall and the confounding effects of water provision and fences

Elephant are increasing across some areas of Africa leading to concerns that they may reduce woodlands through their feeding. Droughts may help limit elephant numbers, but they are generally both episodic and local. To explore more general impacts of rainfall, we examine how its annual variation influences elephant survival across ten sites. These sites span an almost coast-to-coast transect of southern Africa that holds the majority of the ~500,000 remaining savanna elephants. Elephants born in high rainfall years survive better than elephants born in low rainfall years. The relationship is generally weak, except at the two fenced sites, where rainfall greatly influenced juvenile survival. In these two sites, there are also extensive networks of artificial water. Rainfall likely affects elephant survival through its influence on food. The provision of artificial water opens new areas for elephants in the dry season, while fencing restricts their movements in the wet season. We conclude that the combination of these factors makes elephant survival more susceptible to reductions in rainfall. As a result, elephants living in enclosed reserves may be the first populations to feel the impacts of global warming which will decrease average rainfall and increase the frequency of droughts. A way to prevent these elephants from damaging the vegetation within these enclosed parks is for managers to reduce artificial water sources or, whenever practical, to remove fences.     

Drought mortality of bush elephants in Hwange National Park, Zimbabwe

African bush elephants inhabiting the undeveloped Kalahari Sands region of Hwange National Park, Zimbabwe are subject to episodic mortality during droughts. We monitored the drought‐related mortality of elephants in Hwange National Park over the course of an extended drought between 1993 and 1995. The drought‐related mortality of elephants was higher during 1994 than 1995, despite significantly higher rainfall in 1994 than 1995. We found significant differences in the age‐specific mortality of elephants during 1994 and 1995. The cumulative mortality profile from this study differed significantly from previous die‐offs at this site, with a higher mortality among adult age classes than that reported from earlier studies in Hwange National Park. The effective duration of the rainy season, not total annual precipitation, appears to be the best predictor for the potential severity of drought mortality among elephants in the Kalahari Sands habitats of Hwange National Park.     

Bee Threat Elicits Alarm Call in African Elephants

Unlike the smaller and more vulnerable mammals, African elephants have relatively few predators that threaten their survival. The sound of disturbed African honeybees Apis meliffera scutellata causes African elephants Loxodonta africana to retreat and produce warning vocalizations that lead other elephants to join the flight. In our first experiment, audio playbacks of bee sounds induced elephants to retreat and elicited more head-shaking and dusting, reactive behaviors that may prevent bee stings, compared to white noise control playbacks. Most importantly, elephants produced distinctive “rumble” vocalizations in response to bee sounds. These rumbles exhibited an upward shift in the second formant location, which implies active vocal tract modulation, compared to rumbles made in response to white noise playbacks. In a second experiment, audio playbacks of these rumbles produced in response to bees elicited increased headshaking, and further and faster retreat behavior in other elephants, compared to control rumble playbacks with lower second formant frequencies. These responses to the bee rumble stimuli occurred in the absence of any bees or bee sounds. This suggests that these elephant rumbles may function as referential signals, in which a formant frequency shift alerts nearby elephants about an external threat, in this case, the threat of bees.     

How do African elephants utilize the landscape during wet season? A habitat connectivity analysis for Sioma Ngwezi landscape in Zambia

1. The influence of environmental factors on the distribution and persistence of African elephants (Loxodonta africana) is pertinent to policy makers and managers to formulate balanced plans for different land‐use types.
2. The study focuses on movement of elephants and how they utilize foraging areas in Sioma Ngwezi landscape in Zambia by answering the following questions: (1) Which environmental variables and land‐cover class predict the movement of elephants during the wet season in Sioma Ngwezi landscape? (2) What is the wet season suitable habitat for elephants in Sioma Ngwezi landscape? (3) What are the major wet season movement corridors for elephants in Sioma Ngwezi landscape?
3. We used GPS telemetry data from the collared elephants to assess habitat connectivity. Maximum entropy (MaxEnt) and linkage mapper were the tools used to predict habitat suitability, movement corridors, and barriers in the landscape during the wet season.
4. The study identified elevation, land cover, and NDVI as the most important environmental predictors that modify the dispersal of elephants in the landscape during the wet season. Additionally, a total of 36 potential wet season corridors were identified connecting 15 core areas mainly used for foraging and protection from poachers in the landscape. Of these, 24 corridors were highly utilized and are suggested as priority corridors for elephant movement in the landscape.
5. The identified wet season habitats and functional corridors may help to combat elephant poaching by patrolling areas with high relative probability of elephant presence. The findings may also help abate human–elephant conflict such as crop‐raiding by managing identified corridors that run into agriculture zones in the game management area.

     

Ranging behaviour of Uganda’s elephants

Elephant populations are in decline across the African continent, but recent aerial surveys show that populations in Uganda are increasing. However, threats such as poaching and habitat disturbance remain. Having a comprehensive knowledge of the ranging behaviour of Ugandan elephants is crucial to understanding where critical habitat for the species occurs. We investigated various aspects of ranging behaviour of 45 radio-collared elephants (Loxodonta africana) in three areas—Queen Elizabeth Protected Area (QEPA), Murchison Falls (MFPA) Protected Area and Kidepo Valley (KVCA) Conservation Area. We also set Ugandan analyses in a continental context by comparison with home ranges reported in published literature. Elephants within KVCA had larger core ranges than elephants in QEPA or MFPA. Wet season ranges in KVCA were much larger than dry season ranges. The most important core areas in all three national parks were centred around water resources. Home range size was negatively correlated with net primary productivity (NPP) at Ugandan (N = 39 individuals) and continental (N = 17 sites) scales. This study indicates that, at a local scale, factors such as water source location are important in shaping elephant ranging behaviour. At larger scales, factors such as NPP are good predictors of elephant home range size.     

Seasonality of reproduction in Asian elephants Elephas maximus and African elephants Loxodonta africana: underlying photoperiodic cueing?

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Elephants caught in the middle: impacts of war, fences and people on elephant distribution and abundance in the Caprivi Strip, Namibia

We conducted wet [26 March–4 April 2003 (Apr03)] and dry [1–8 November 2005 (Nov05)] season aerial surveys of African elephants ( Loxodonta africana Blumenbach) in the Caprivi Strip, Namibia to provide an updated status report on elephant numbers and distribution and assist with a historical analysis of elephant distribution and abundance in the Caprivi Strip. During the wet season when water was available in seasonal pans, elephants were widely distributed throughout the survey area. In contrast, during the dry season, a majority of elephant herds occurred within 30 km of the perennial Kwando, Linyanti and Okavango rivers and few herds occurred within the West Caprivi Game Reserve where water in the seasonal pans was limited. We estimated 5318 elephants for the 7731-km² survey area (0.71 elephants km⁻²) for the Apr03 wet season survey and 6474 elephants for the 8597-km² survey area (0.75 elephants km⁻²) for the Nov05 dry season survey. Based on our aerial surveys and reports of elephant numbers and distribution from historical aerial surveys and telemetry studies, civil war, veterinary fences and human activities appear to have effected changes in African elephant abundance, distribution and movements in the Caprivi Strip, Namibia since 1988 when the first comprehensive aerial surveys were conducted.     

Elephant Genomes Reveal Accelerated Evolution in Mechanisms Underlying Disease Defenses

Disease susceptibility and resistance are important factors for the conservation of endangered species, including elephants. We analyzed pathology data from 26 zoos and report that Asian elephants have increased neoplasia and malignancy prevalence compared with African bush elephants. This is consistent with observed higher susceptibility to tuberculosis and elephant endotheliotropic herpesvirus (EEHV) in Asian elephants. To investigate genetic mechanisms underlying disease resistance, including differential responses between species, among other elephant traits, we sequenced multiple elephant genomes. We report a draft assembly for an Asian elephant, and defined 862 and 1,017 conserved potential regulatory elements in Asian and African bush elephants, respectively. In the genomes of both elephant species, conserved elements were significantly enriched with genes differentially expressed between the species. In Asian elephants, these putative regulatory regions were involved in immunity pathways including tumor-necrosis factor, which plays an important role in EEHV response. Genomic sequences of African bush, forest, and Asian elephant genomes revealed extensive sequence conservation at TP53 retrogene loci across three species, which may be related to TP53 functionality in elephant cancer resistance. Positive selection scans revealed outlier genes related to additional elephant traits. Our study suggests that gene regulation plays an important role in the differential inflammatory response of Asian and African elephants, leading to increased infectious disease and cancer susceptibility in Asian elephants. These genomic discoveries can inform future functional and translational studies aimed at identifying effective treatment approaches for ill elephants, which may improve conservation.     

Elephant spatial use in wet and dry savannas of southern Africa

The influence of elephants on woody vegetation cover varies from place to place. In part this may be due to the way elephants utilize space across landscapes and within their home ranges in response to the availability and distribution of food. We used location data from 18 cows at six study sites across an east to west rainfall gradient in southern Africa to test whether wet- and dry-season home-range sizes, evenness of space use within seasonal home ranges and range overlap between seasons and between years, differed between wet and dry savannas. We then tested whether the quantity, distribution and seasonal stability in vegetation productivity, a coarse measure of food for elephants, explained differences. Elephants in wet savannas had smaller wet- and dry-season home ranges and also returned to a higher proportion of previously visited grid cells between seasons and between years than elephants living in dry savannas. Wet-season home-range sizes were explained by seasonal vegetation productivity while dry-season home-range sizes were explained by heterogeneity in the distribution of vegetation productivity. The influence of the latter on dry-season home ranges differed among structural vegetation classes. Range overlap between seasons and between years was related to inter-seasonal and inter-annual stability in vegetation productivity, respectively. Evenness of elephant spatial use within home ranges did not differ between savanna types, but it was explained by seasonal vegetation productivity and heterogeneity in the distribution of vegetation productivity during the wet season. Differences in elephant spatial use patterns between wet and dry savannas according to vegetation structure and season may need to be included in the development of site-specific objectives and management approaches for African elephants.     

Sexual segregation in habitat use by elephants in Chobe National Park, Botswana

We report on a study conducted on free‐ranging African elephants in the woodlands of northern Botswana. We compared bull groups and family units with regard to (1) their patterns of habitat use and (2) their ranging distances from perennial water sources. During the dry season, adult males frequented more habitat types than family units, whereas family units used a wider diversity of habitats than bulls during the wet season. Bulls roamed widely (>10 km) from perennial drinking water in the dry season, when family units congregated within 3.5 km of the rivers. During the wet season, when ephemeral pans were abundant, all elephant groups were found at intermediate distances (5 km) from the rivers. The spacing of elephants in the dry season is consistent with sexual segregation but we reject the hypothesis that this is an outcome of indirect competition for food, because our concurrent studies on elephant feeding ecology found no evidence for intraspecific competition. Instead, we propose that most adult male elephants space themselves to avoid conflict with musth bulls and roam widely in the dry season between discretely distributed feeding ‘hotspots’. The small proportion of males that are in musth remain close to family units to maximize mating opportunities, and family units are unable to range far from water in the dry season. This is due to (1) comparatively high rates of water turn‐over among juveniles and lactating cows and (2) the reduced mobility of neonates.     

Elephant distribution around a volcanic shield dominated by a mosaic of forest and savanna (Marsabit, Kenya)

We investigated the factors that influenced the distribution of the African elephant around a volcanic shield dominated by a mosaic of forest and savanna in northern Kenya. Data on elephant distribution were acquired from four female and five bull elephants, collared with satellite-linked geographical positioning system collars. Based on the eigenvalues (variances) of the correlation matrix, the six factors that contributed significantly to high total variances were distance from drinking water (24%), elevation (15%), shrubland (10%), forest (9%), distance from settlements (8%) and distance from minor roads (7%), contributing to 73% in the observed variation of the elephant distribution. The elephants were found at high forested elevations during the dry season but they moved to the lowlands characterized by shrubland during the wet season. Elevation acts as a proxy for the vegetation structure. The presence of elephants near permanent water points (13%) and seasonal rivers (11%) during the dry and wet seasons, respectively, demonstrates that water is the most important determinant of their distribution throughout the year. We conclude that the distribution of elephants in Marsabit Protected Area and its adjacent areas is influenced mainly by drinking water and vegetation structure.     

Observations on the length of the intestinal tract of African <Emphasis Type="Italic">Loxodonta africana</Emphasis> (Blumenbach 1797) and Asian elephants <Emphasis Type="Italic">Elephas maximus</Emphasis> (Linné 1735)

The digestive tract of elephants is surprisingly short compared to other herbivorous mammals. However, measurements relating the length of the intestine to the body mass of the respective individual are rare. In this study, we report such data for an African elephant and an Asian elephant. Our data support the hypothesis that Asian elephants have a longer intestinal tract than their African counterparts. These findings are in accord with the observation of longer retention times and higher digestion coefficients in Asian as compared to African elephants. This difference between the species could be the reflection of slightly different ecological niches, with Asian elephants adapted to a natural diet with a higher proportion of grass.     

Is there a future for elephants in West Africa?

The elephants of West Africa have experienced a long history of human disturbance. Before 1800 they were much affected by the precolonial empires of the savanna and Sahelian zones, the trans-Saharan trade routes, and the coastal trade established by the Europeans. During the 19th century, the increasing demand for ivory from Europe and North America, the European penetration of the region, and the evolution of breech-loading rifles devastated the remaining elephants. The elephant population of West Africa collapsed before the outbreak of World War I because of intense hunting for ivory. This collapse pre-empted the decline that would have occurred anyway due to the rapid growth of the human population and consequent loss of habitat. Elephants now find themselves in about 70 small isolated populations that cover only 5% of the region. These fragments are very vulnerable – whether in the arid lands or the humid forests – to poaching and general human disturbance. There are few data on numbers; most of the population estimates are guesses. Two-thirds of the populations are thought to consist of fewer than 200 animals and therefore have a low probability of surviving the next century. As more habitat is lost to human activities, West African elephants will soon remain only in protected areas. But many parks and reserves are managed poorly and cannot offer effective protection; they do not guarantee a future for elephants. In addition, their crop-raiding habit makes elephants unpopular in rural communities surrounding protected areas. Human populations are expected to continue growing and resources for conservation are scarce. The future of West African elephants lies in a small network of well-protected areas.     

Seed dispersal kernel of the largest surviving megaherbivore—the African savanna elephant

Owing to the late Pleistocene extinctions, the megafauna of Europe, Australia and the Americas disappeared, and with them the dispersal service they offered megafaunal fruit. The African savanna elephant, the largest remaining megaherbivore, offers valuable insights into the seed dispersal services provided by extinct megafauna in prehistoric times. Elephant seed dispersal studies have for the most part concentrated on African and Asian forest elephants. African savanna elephants are morphologically distinct from their forest counterparts. Like the forest elephants they consume large quantities of fruit from a large number of tree species. Despite this little is known of the savanna trees that rely on elephants for their dispersal or the spatial scale at which these seeds are dispersed. We combined information from feeding trials conducted on four park elephants with field telemetry data from 38 collared elephants collected over an 8‐year period in APNR/Kruger National Park to assess the seed dispersal service provided by savanna elephants. This study provides the first detailed account of the spatial scale at which African savanna elephants disperse seeds. Our mechanistic model predicts that 50 percent of seeds are carried over 2.5 km, and distances up to 65 km are achievable in maximum gut passage time. These findings suggest the savanna elephant as the longest distance terrestrial vertebrate disperser yet investigated. Maintaining their ecological role as a seed disperser may prove a significant factor in the conservation of large‐fruited tree diversity within the savannas. These results suggest that extinct megafauna offered a functionally unique dispersal service to megafaunal fruit.     

Determining adrenocortical activity as a measure of stress in African elephants (Loxodonta africana) in relation to human activities in Serengeti ecosystem

African elephants (Loxodonta africana) play a vital role in most African ecosystems, with their opportunity to alter the entire ecosystem by their sheer numbers. Defining and measuring animal welfare has been much discussed. One potential way of determining an animal's welfare is to record the absence or presence of stress. Little research on elephant welfare has so far been performed in the Serengeti ecosystem. The aim of this study was to record the faecal glucocorticoid metabolite levels of African elephants in areas with high or with minimum human interference. A total of 117 faecal samples were collected from randomly located single elephants as well as family herds in the northern, central and western Serengeti National Park (SNP) as well as in Grumeti Game Reserve and Ikoma Open Area, northern Tanzania in 2010. Elephants had higher levels of faecal glucocorticoid metabolites in the areas outside, compared with areas inside SNP. No single males were observed outside SNP, and in general, higher abundance of elephants was observed inside SNP. This suggests that elephants may prefer to reside in the potential safer areas inside the national park, demonstrating the importance of protected areas to improve the welfare of elephants.     

The historical development of zoo elephant survivorship

In the discussion about zoo elephant husbandry, the report of Clubb et al. (2008, Science 322: 1649) that zoo elephants had a “compromised survivorship” compared to certain non-zoo populations is a grave argument, and was possibly one of the triggers of a large variety of investigations into zoo elephant welfare, and changes in zoo elephant management. A side observation of that report was that whereas survivorship in African elephants (Loxodonta africana) improved since 1960, this was not the case in Asian elephants (Elephas maximus). We used historical data (based on the Species360 database) to revisit this aspect, including recent developments since 2008. Assessing the North American and European populations from 1910 until today, there were significant improvements of adult (≥10 years) survivorship in both species. For the period from 1960 until today, survivorship improvement was significant for African elephants and close to a significant improvement in Asian elephants; Asian elephants generally had a higher survivorship than Africans. Juvenile (<10 years) survivorship did not change significantly since 1960 and was higher in African elephants, most likely due to the effect of elephant herpes virus on Asian elephants. Current zoo elephant survivorship is higher than some, and lower than some other non-zoo populations. We discuss that in our view, the shape of the survivorship curve, and its change over time, are more relevant than comparisons with specific populations. Zoo elephant survivorship should be monitored continuously, and the expectation of a continuous trend towards improvement should be met.     

Differences in woody vegetation are unrelated to use by African elephants (Loxodonta africana) in Mkhuze Game Reserve, South Africa

The reintroduction of African elephants into fenced game reserves throughout South Africa has presented managers with several challenges. Although elephants are a natural part of southern African ecosystems, their confinement to fenced protected areas in South Africa has exacerbated their potential to impact their habitats negatively. However, many studies investigating the impact of elephants have failed to control for the effects of other browsers on the vegetative community. In this study, we used location data on an elephant herd to delineate high-use and low-use areas. This paired design allowed us to minimize confounding factors that could explain differences in the structure, diversity and utilization of woody species. We found little evidence to suggest elephant-mediated change in, or selection for, the structure or diversity of woody species; however, our results suggest that elephants may be altering the composition of species by preferentially using areas with higher canopy diversity and by enhancing sapling recruitment. Although stripping of bark was higher in high-use areas, there was no evidence of differential mortality of tree species. Therefore, in our study area, and over the current time scale, elephants are having a negligible impact on the vegetative community.     

Do elephants prevent other African herbivores from using waterholes in the dry season?

In some African protected areas, concerns have arisen about the influence of locally high elephant numbers on other forms of biodiversity. In arid and semi-arid savannas, surface-water resources are scarce and agonistic interactions between elephants and other herbivores have been reported at waterholes, yet surprisingly very little is known about the impact of elephants on the use of waterholes by other herbivores. Here, we test whether when there are elephants at a waterhole, other herbivores (1) do not change their drinking behaviour; (2) spend shorter time around the water because they are disturbed by elephants’ presence and consequently have to leave the waterhole area probably without having met their water requirements, or (3) spend more time around the water probably owing to an increase in vigilance activities or because the presence of elephants may signal safety from predators. Results show that all species spend longer time around water when there are elephants at the waterhole, although the difference is not large. Consequently, this study strongly suggests that elephants do not prevent other herbivores from drinking (time at waterholes is not shortened when elephants are around). Further, if the additional time spent to drink is linked to an increased vigilance, the difference is not large, and hence unlikely to affect the population dynamics of other herbivores.     

Ovarian function in the elephant: luteinizing hormone and progesterone cycles in African and Asian elephants

Serum samples were collected weekly for 3 yr from two female African elephants, for 18 mo from two other female African elephants, and for 2 yr from two female Asian elephants. Animals were not sedated at the time of blood collection. Ovarian cycles, characterized by changes in progesterone and immunoreactive luteinizing hormone (ILH) concentrations, averaged 15.9 +/- 0.6 wk (N = 25) for African females and 14.7 +/- 0.5 wk for Asian females (N = 10). The length of the active luteal phase averaged 10.0 +/- 0.3 wk for African elephants (range 8-14 wk) and 10.6 +/- 0.6 wk for Asian females (range 9-13 wk). Interluteal phases were 5.9 +/- 0.6 wk for African females and 4.2 +/- 0.5 wk for Asian females. One African female (Maliaca) had two extended interluteal phases, both occurring between the months of February and May. Excluding these two periods, there were no differences in the length of the ovarian cycle or the length of the luteal phase between species of elephant. Serum progesterone in both species ranged from less than 50 pg/ml to 933 pg/ml. Average progesterone concentrations during the luteal phase were significantly lower in African elephants compared with Asian elephants (328 +/- 13, N = 30 cycles vs. 456 +/- 23, N = 14 cycles; p less than 0.001). ILH ranged from nondetectable to 11.6 ng/ml. These data suggest that the length of the ovarian cycle in the African elephant is about 16 wk and confirm that the length of the ovarian cycle in the Asian elephant is about 15 wk.     

Exploitation of underground bee nests by three sympatric consumers in Loango National Park,Gabon

Honey represents a highly nutritious resource for animals, but is difficult to obtain given bees' defensive strategies. We investigated exploitation of the underground nests of stingless bees (Meliplebeia lendliana) by three sympatric consumers in Loango National Park, Gabon: the central African chimpanzee (Pan troglodytes troglodytes), forest elephant (Loxodonta cyclotis) and honey badger (Mellivora capensis). Given the differences in their respective morphological traits and sensory abilities, we hypothesized that chimpanzees would be more limited in digging out the bee nests, compared to the other two competitors, and would show behavioral strategies to overcome such constraints. Our dataset comprised camera trap footage recorded over 60 mo at 100 different bee nests. Chimpanzees visited the nests more often than the other consumers, showing a frequency of extraction success comparable to that observed in honey badgers, the most efficient digger. Both chimpanzees and honey badgers increased their extractive attempts across the dry season, whereas elephants did not. The soil hardness was greater during the dry season than the wet season and, possibly in order to compensate for this, chimpanzees showed a tendency toward digging at nests found in relatively softer soil. They also seemed to be inhibited by indirect cues left by other consumers, possibly as a risk‐avoidance strategy. Overall, chimpanzees and honey badgers extracted the underground nests of stingless bees with similar frequencies, whilst forest elephants did so only occasionally. Moreover, chimpanzees can use tools and other behavioral strategies to overcome the physical limitations that may constrain their exploitation of this resource.