Sacoglottis gabonensis– a keystone fruit for forest elephants in the Réserve de Faune du Petit Loango, Gabon

This study suggests that the fruits of Sacoglottis gabonensis (Baill.) Urb. (Humiriaceae) are a keystone resource for forest elephants ( Loxodonta cyclotis Matschie) in a coastal rain forest, the Réserve de Faune du Petit Loango, Gabon (now part of Loango National Park). Faecal counts demonstrated that forest elephants used Sacoglottis -dominated forest more when Sacoglottis was abundant and electivity indices suggest that Sacoglottis is a preferred food. The flora of Petit Loango is characterized by the absence of herbaceous vegetation such as Marantaceae and Zingiberaceae, and during the prolonged dry season few fleshy fruits are present other than Sacoglottis fruits, which are produced in a glut during this time. While inter-annual fruiting reliability remains to be confirmed, fruit production in 1998 and high stem density relative to other study sites provide indirect evidence that Sacoglottis fruits are a reliable inter-annual resource at Petit Loango. It is thus proposed that Sacoglottis gabonensis fruits fulfil an important role as a keystone 'fallback' resource for forest elephants during the dry season at Petit Loango.     

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.     

Relatedness and demography of African forest elephants: inferences from noninvasive fecal DNA analyses

African forest elephants (Loxodonta cyclotis) are genetically and morphologically distinct from their savannah counterparts, but their biology remains poorly understood. In this study, I use noninvasive fecal DNA analyses to examine the relatedness structure and historical demography of forest elephants at 2 sites in SW Gabon, central Africa. Pairwise relatedness values calculated between 162 elephant individuals genotyped at 8 microsatellite loci were significantly higher within spatially associated dung piles than between random pairings for one site. First- and second-order relatives were most commonly detected among dung piles from adult female pairs and adult females and juveniles. Pairwise relatedness estimates suggested that, like savannah elephants, forest groups are largely composed of adult females, their sisters, and juvenile offspring. Associations between males, and groups containing juveniles from multiple related females, were detected but at much lower frequency. Analysis of mitochondrial d-loop sequences from 70 elephant individuals identified 2 haplogroups in SW Gabon.     

Physiological stress and refuge behavior by African elephants

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

Small-scale dung survey reveals high forest elephant density and preference for mixed species forest in an intact protected area

Understanding the ecological factors influencing African forest elephant (Loxodonta cyclotis) abundance and distribution is crucial for their conservation management in Central Africa. Dung surveys have been conducted at the landscape scale and confirmed the overwhelming impact of anthropogenic activities on forest elephants. We present results from a small-scale survey in a pristine protected area without anthropogenic activities to elucidate the ecological factors influencing forest elephant density. We conducted a line transect dung survey in a small study area (110 km²) around Mbeli Bai, a natural forest clearing in the Nouabalé-Ndoki National Park, Republic of Congo, and compare results with a landscape survey conducted during the same period. We used habitat specific dung decay data collected on site to estimate elephant density using distance sampling. We fitted Generalized Additive Models to elephant dung encounter rate using explanatory variables collected during the transect survey and from geospatial data. The small-scale survey revealed a precise estimate of forest elephant density that was twice as high as the result from a landscape survey with higher density in mixed species forest for the small-scale survey. We could not find an impact of the proximity of forest clearings and proximity to rivers at the small scale. Fine-scale habitat features, e.g. degree of canopy and understorey closure, had little explanatory power for elephant dung encounter rate. Small-scale dung surveys are a useful method to reveal spatio-temporal variation in forest elephant density and distribution which can inform conservation practitioners in a timely manner. Combining monitoring methods at various spatial scales improves our knowledge and conservation efforts of forest elephants. Nouabalé-Ndoki National Park is a stronghold for forest elephants and of global importance for their conservation.

     

Assessing the effect of roads on impala (Aepyceros melampus) stress levels using faecal glucocorticoid metabolites

In Tanzania, a proposed road through the Serengeti National Park has caused international debate as to whether it will cause irreversible damage to the ecosystem. We assessed the effect of current roads and traffic in the Serengeti ecosystem on a model ungulate species, the impala antelope (Aepyceros melampus). We measured how faecal glucocorticoid metabolites (FGM), an indicator of physiological stress levels, were affected by different levels of traffic while accounting for other stress‐related variables such as distance to roads, vegetation type and group size. A total of 196 faecal samples were collected. FGM were measured using an enzyme immunoassay (EIA) validated for several ruminants. FGM levels were significantly elevated near major roads with higher traffic levels compared to roads with less traffic. Predators and illegal hunting might also affect FGM levels in the impala, but the potential effect of these variables was inconsistent with the recorded FGM levels and cannot explain the observed variation. These results show for the first time that an African mammal exhibits significant physiological stress in relation to roads and traffic.     

Group size, density and biomass of large mammals in the Réserve de Faune du Petit Loango, Gabon

Group size, density and biomass of large‐bodied diurnal mammal species in the Réserve de Faune du Petit Loango, Gabon (now Parc National de Loango) was determined over a 12‐month period using standard line‐transect methods. Petit Loango encompasses a range of distinct habitat types, including coastal scrub, savanna, swamps and disturbed and mature forest. Such intact coastal habitats are increasingly rare on the Central‐West African coastline. Faecal censusing indicated highest forest elephant (Loxodonta africana cyclotis) and buffalo (Syncerus caffer nanus) ecological densities at the extreme coast (2.48 and 1.29 km⁻² respectively), probably reflecting high intensity of use of this habitat. Ape density was comparable with that at other Central African study sites at 1.01 individuals km⁻². Mean total biomass of diurnal primates, elephants and other ungulates over the 20 km² site was 3290 kg km⁻². Forest elephants and red river hogs (Potamochoerus porcus) constituted the bulk of the biomass, at 67% and 14% respectively. Primates made up 5% of the biomass. This is the first estimation of mammal density and biomass over an annual cycle at a Central African coastal site, and provides baseline data for long‐term studies in such habitats and to aid habitat and wildlife management decisions.     

Endocrine Correlates of Musth in Free-Ranging Asian Elephants (Elephas maximus) Determined by Non-Invasive Faecal Steroid Hormone Metabolite Measurements

The occurrence of musth, a period of elevated levels of androgens and heightened sexual activity, has been well documented for the male Asian elephant (Elephas maximus). However, the relationship between androgen-dependent musth and adrenocortical function in this species is unclear. The current study is the first assessment of testicular and adrenocortical function in free-ranging male Asian elephants by measuring levels of testosterone (androgen) and cortisol (glucocorticoid – a physiological indicator of stress) metabolites in faeces. During musth, males expectedly showed significant elevation in faecal testosterone metabolite levels. Interestingly, glucocorticoid metabolite concentrations remained unchanged between musth and non-musth periods. This observation is contrary to that observed with wild and captive African elephant bulls and captive Asian bull elephants. Our results show that musth may not necessarily represent a stressful condition in free-ranging male Asian elephants.     

Stress Response of Working African Elephants to Transportation and Safari Adventures

Abstract: African elephants (Loxodonta africana) are intensively managed in southern Africa and are routinely translocated between reserves. Domesticated elephants are used for elephant-back safaris and interactions with guests. Understanding how elephants respond to such activities is critical because of welfare issues associated with both humans and elephants. We investigated the stress response (i.e., fecal glucocorticoid metabolite secretion [FGM]) of working elephants in Letsatsing Game Reserve, South Africa, over 1 year to evaluate their response to transportation and ecotourism activities. We used free-ranging elephants in adjacent Pilanesburg National Park as controls. Fecal glucocorticoid metabolites were greatest prior to and during translocation and declined over the year. Within 1–2 months of transportation, FGM levels in working elephants became indistinguishable from those in wild elephants. Fecal glucocorticoid metabolite levels were higher during human interaction days than days without interaction. The highest observed FGM levels were associated with transportation and episodic loud noises. Transportation is a stressful activity for elephants, and ≥3 months should be provided to translocated elephants to acclimate to their new surroundings. Although stress levels of elephants increased slightly when interacting with humans in the contexts we studied, evaluating interactions under a wider range of contexts is necessary to minimize danger to elephants and humans.     

Diet and distribution of elephant in the Maputo Elephant Reserve, Mozambique

The distribution and diet of the elephants of the Maputo Elephant Reserve were studied using dung counts, satellite tracking and faecal analysis. The results were compared with earlier data from before the civil war in Mozambique. The elephant population decreased during the civil war, but 180 animals still remain. Earlier studies described the elephants as preferring the grass plains. Currently, the elephants prefer the dense forest patches over the high quality forage found in the grass plains. Water salinity affected distribution; elephant dung piles were found closer to fresh water in the dry season. A total of 95 different plant species were identified in the faeces. The percentage of grass was relatively low compared with other studies, increasing at the beginning of the rainy season. At the end of the dry season, elephants concentrated on the few available browse species with young leaves, but generally preferred grass species to browse species. Diet composition was mainly affected by season and less by habitat. The elephants have changed their habitat preference in reaction to poaching, and probably increased the contribution of browse species in the diet. The presence of forest patches has been vital for the survival of the elephants.     

Forest elephant group composition, frugivory and coastal use in the Réserve de Faune du Petit Loango, Gabon

Results of a 16‐month study of forest elephant (Loxodonta africana cyclotis Matschie) group size and composition, coastal use and frugivory are presented from the Réserve de Faune du Petit Loango, Gabon (now the Parc National de Loango). Mean forest elephant group size was 2.2 (n = 140) including solitary animals, and 3.1 excluding solitaries. Elephants consumed fruits of at least 49 species, and of the 220 elephant dungpiles examined, 185 (84.1%) contained seeds and 203 (92.3%) contained the remains of fruits (seeds and/or pulp). The mean number of seed species per dungpile (±SD) was 2.01 ± 1.49, and the mean number of fruit species was 2.28 ± 1.43. Elephants used the coastal habitat more during warmer months, and during the afternoon than the morning. It is hypothesized that coastal habitat use is related to sodium intake through consumption of salt‐coated vegetation.     

Forest elephant paths and movements at the Odzala National Park, Congo: the role of clearings and Marantaceae forests

The role of forest clearings and Marantaceae forest on elephant movement at the Odzala National Park was studied by describing and mapping elephant paths. Three types of paths were distinguished based on their direction, length, width, elephant activity and the types of forests they cross. They were called boulevards, foraging paths, and clearing alleys. All paths explored were heavily used. Boulevards are used for travelling over long distances (up to 34 km) and for linking up rapidly favourite sites such as clearings. Foraging paths mainly run through the medium-density Marantaceae forests which provide elephants with both herbaceous foods and tree fruit. Clearing alleys form a dense network of tracks around clearings. This network could be a result of an anti-predatory monitoring behaviour of elephants before entering the clearings where they have long experienced heavy poaching. The heavy trampling which results from the elephant activity could contribute to the maintenance of clearings by preventing germination and seedling survival of the numerous seeds dispersed in their dung.     

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.     

Estimating population sizes for elusive animals: the forest elephants of Kakum National Park,Ghana

African forest elephants are difficult to observe in the dense vegetation, and previous studies have relied upon indirect methods to estimate population sizes. Using multilocus genotyping of noninvasively collected samples, we performed a genetic survey of the forest elephant population at Kakum National Park, Ghana. We estimated population size, sex ratio and genetic variability from our data, then combined this information with field observations to divide the population into age groups. Our population size estimate was very close to that obtained using dung counts, the most commonly used indirect method of estimating the population sizes of forest elephant populations. As their habitat is fragmented by expanding human populations, management will be increasingly important to the persistence of forest elephant populations. The data that can be obtained from noninvasively collected samples will help managers plan for the conservation of this keystone species.     

Estimating forest elephant density in Sapo National Park (Liberia) with a rainfall model

The number of elephant dung‐piles lying on the forest floor is a function of the number of elephants present and the rainfall in the 2 preceding months. We present the results of a stochastic model that describes this relationship and we show how it can be used to estimate elephant numbers. The data from a survey in Sapo NP (Liberia) in 1989 are used as an example. The dung‐pile density was estimated at 152 km^?2 with confidence interval from 72 to 322, and the number of elephants was estimated to be 313 with confidence interval from 172 to 617.     

Population structure and behaviour of crop-raiding elephants in Kibale National Park, Uganda

We examined patterns of crop raiding by elephants across gender and age classes in relation to elephant life history and sociobiology and estimated the quantitative contribution of crops to elephant diet in Kibale National Park (KNP). Elephant dung‐boli sizes were used to estimate age and sex, while the presence of crop remains in the dung of crop‐raiding elephants was used as evidence of repeated raiding. To estimate the expected proportion of elephants raiding per age class, the age distribution of raiders was compared with the age distribution of all KNP elephants. Elephants raiding crops were predominantly males. They began raiding in expected proportions at 10–14 years while a higher than expected proportion raided crops at 20–24 years. These results suggest that crop raiding is initiated at an age when male elephants leave their families and a large proportion of elephants raid when they are approaching reproductive competition. Evidence from dung of crop raiders, suggests that repeated raiding increases with age. Crop raiders derived 38% of their daily forage from the short time spent raiding, consistent with expectations of foraging theory. Males may be more likely to learn crop raiding because they are socially more independent and experience intense mating competition than females.     

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

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

Working with mahouts to explore the diet of work elephants in Myanmar (Burma)

At an elephant camp in central Myanmar (Burma), we interviewed mahouts and veterinarians to describe the diet of Asian elephants (Elephas maximus) in a mixed-deciduous forest. Elephants showed a broad dietary breadth (103 plant species from 42 families); consumed mostly browse (94% of plant species); and were very selective about plant parts [e.g., many trees were eaten exclusively for their bark (22%) or fruits (14%)]. The fruits from 29 plant species were recorded to be eaten by elephants. Several of these were found as fruit remains, seeds, or seedlings in elephant dung, suggesting a role of Asian elephants in seed dispersal. Work elephants and their mahouts prove to be a rich source of information to understand wild elephant ecology.     

Seed dispersal of Acacia erioloba by African bush elephants in Hwange National Park, Zimbabwe

Approximately two-thirds (64%) of all dry season samples of elephant dung analysed during a 3-year study in the Main Camp subregion of Hwange National Park, Zimbabwe, contained seed and/or pod materials from Acacia erioloba . Most seeds were recovered intact and actively germinating seeds were not uncommon. Very little pod mass relative to seed mass was recovered in most instances, with pod fragments recorded from only 56% of all exhaustively sampled elephant dung piles containing A. erioloba fruit materials. Nonetheless, large pod fragments and even entire intact pods were recovered occasionally from elephant dung. Seeds and pods of A. erioloba may comprise 12% or more of total wet-weight dung mass; individual dung piles were found which contained > 5000 A. erioloba seeds. Birds and smaller mammals search out and consume A. erioloba seeds present within elephant dung piles. The findings of this study indicate that potential digestibility of A. erioloba seeds for bush elephants ( Loxodonta africana africana ) may be much higher than expected from previous studies. In controlled feeding trials with captive bush elephants (age 11–15 years old) maintained on predominantly free-range dry season diets, the estimated efficiency of digestion for A. erioloba seeds consumed in pods was 81% to 96%, with a gut-transit time of between 24.5 and 36.0 h. On the basis of throughput times determined in experimental feeding trials, potential elephant-dispersal distances of 20–50 km are predicted for A. erioloba in the Kalahari Sands landscapes of southern central Africa.     

Dry season habitat selection by a recolonizing population of Asian elephants<Emphasis Type="Italic">Elephas maximus</Emphasis> in lowland Nepal

Owing to landclearing and human expansion, Asian elephantElephas maximus Linnaeus, 1758 is declining throughout its range. In lowland Nepal, the species now only occurs in small remnant populations, shared with India. In order to develop guidelines for conserving the species in the country, we studied the habitat use of a small and recently re-established population in Bardia National Park. We used the distribution of dung in fixed width transects to estimate seasonal habitat selection at a general scale of the Park. We also analyzed a specific habitat selection by elephants within the sal-dominated forest, by comparing the composition of trees and frequency of previous elephant impact on them along fresh tracks with those at random points. Elephants strongly preferred floodplain communities both during the cool and the hot season, but there was a marked shift from forest to grass-dominated subtypes between these seasons. Within the sal-dominated forest, there were more trees with previous elephant impact and a higher density of important food trees, especiallyMallotus phillippinensis along fresh tracks than in random points. We found little if any effect of human activity or location of available water on the spatial distribution of elephant dung. The density of the colonizing population was low (ca 0.2 animals/km²), but numbers are expected to increase in the future. With the preferred floodplain habitat being quite small (ca 60 km²), animals are then expected to spread outside the national park. A large tract of government forest adjacent to the park may then, for some time, provide needed space for the expanding population.