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.     

Devastating Decline of Forest Elephants in Central Africa

African forest elephants– taxonomically and functionally unique–are being poached at accelerating rates, but we lack range-wide information on the repercussions. Analysis of the largest survey dataset ever assembled for forest elephants (80 foot-surveys; covering 13,000 km; 91,600 person-days of fieldwork) revealed that population size declined by ca. 62% between 2002–2011, and the taxon lost 30% of its geographical range. The population is now less than 10% of its potential size, occupying less than 25% of its potential range. High human population density, hunting intensity, absence of law enforcement, poor governance, and proximity to expanding infrastructure are the strongest predictors of decline. To save the remaining African forest elephants, illegal poaching for ivory and encroachment into core elephant habitat must be stopped. In addition, the international demand for ivory, which fuels illegal trade, must be dramatically reduced.     

Reconciling apparent conflicts between mitochondrial and nuclear phylogenies in African elephants

Conservation strategies for African elephants would be advanced by resolution of conflicting claims that they comprise one, two, three or four taxonomic groups, and by development of genetic markers that establish more incisively the provenance of confiscated ivory. We addressed these related issues by genotyping 555 elephants from across Africa with microsatellite markers, developing a method to identify those loci most effective at geographic assignment of elephants (or their ivory), and conducting novel analyses of continent-wide datasets of mitochondrial DNA. Results showed that nuclear genetic diversity was partitioned into two clusters, corresponding to African forest elephants (99.5% Cluster-1) and African savanna elephants (99.4% Cluster-2). Hybrid individuals were rare. In a comparison of basal forest "F" and savanna "S" mtDNA clade distributions to nuclear DNA partitions, forest elephant nuclear genotypes occurred only in populations in which S clade mtDNA was absent, suggesting that nuclear partitioning corresponds to the presence or absence of S clade mtDNA. We reanalyzed African elephant mtDNA sequences from 81 locales spanning the continent and discovered that S clade mtDNA was completely absent among elephants at all 30 sampled tropical forest locales. The distribution of savanna nuclear DNA and S clade mtDNA corresponded closely to range boundaries traditionally ascribed to the savanna elephant species based on habitat and morphology. Further, a reanalysis of nuclear genetic assignment results suggested that West African elephants do not comprise a distinct third species. Finally, we show that some DNA markers will be more useful than others for determining the geographic origins of illegal ivory. These findings resolve the apparent incongruence between mtDNA and nuclear genetic patterns that has confounded the taxonomy of African elephants, affirm the limitations of using mtDNA patterns to infer elephant systematics or population structure, and strongly support the existence of two elephant species in Africa.     

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.     

Using presence‐only modelling to predict Asian elephant habitat use in a tropical forest landscape: implications for conservation

Aim Asian elephants, Elephas maximus, are threatened throughout their range by a combination of logging, large scale forest conversion and conflict with humans. We investigate which environmental factors, both biotic and abiotic, constrain the current distribution of elephants. A spatially explicit habitat model is constructed to find core areas for conservation and to assess current threats. Location Ulu Masen Ecosystem in the province of Nanggroe Aceh Darussalam on the island of Sumatra, Indonesia. Methods A stratified survey was conducted at 12 sites (300 transects) to establish the presence of elephants. Presence records formed the basis to model potential habitat use. Ecological niche factor analysis (ENFA) is used to describe their niche and to identify key factors shaping elephant distribution. An initial niche model was constructed to describe elephant niche structure, and a second model focused on identifying core areas only. To assess the threat of habitat encroachment, overlap between the elephants’ optimal niche and the occurrence of forest encroachment is computed. Results Elephants were recorded throughout the study area from sea level to 1600 m a.s.l. The results show that the elephant niche and consequently habitat use markedly deviates from the available environment. Elephant presence was positively related to forest cover and vegetation productivity, and elephants were largely confined to valleys. A spatially explicit model showed that elephants mainly utilize forest edges. Forest encroachment occurs throughout the elephants range and was found within 80% of the elephants’ ecological niche. Main conclusions In contrast to general opinion, elephant distribution proved to be weakly constrained by altitude, possibly because of movement routes running through mountainous areas. Elephants were often found to occupy habitat patches in and near human‐dominated areas. This pattern is believed to reflect the displacement of elephants from their former habitat.     

Foraging impacts of Asian megafauna on tropical rain forest structure and biodiversity

Megaherbivores are known to influence the structure, composition, and diversity of vegetation. In Central Africa, forest elephants act as ecological filters by breaking tree saplings and stripping them of foliage. Much less is known about impacts of megafauna on Southeast Asian rain forests. Here, we ask whether herbivory by Asian megafauna has impacts analogous to those of African forest elephants. To answer this, we studied forest (1) structure, (2) composition, (3) diversity, and (4) tree scars in Belum and Krau, two protected areas of Peninsular Malaysia, and compared the results with those obtained in African forests. Elephants are abundant in Belum but have been absent in Krau since 1993. We found that stem density and diversity, especially of tree saplings, were higher in Krau than in Belum. Palms and other monocots were also more abundant in Krau. In Belum, however, small monocots (<1 m tall) were very abundant but larger ones (>1 m tall) were virtually absent, suggesting size‐selective removal. The frequency of stem‐break scars was equal at Belum and Krau but less than in Central Africa and greater than in the Peruvian Amazon where tapirs are the only megafauna. Pigs and tapirs could also contribute to the high frequency of tree scars recorded in Malaysian forests. Forest‐dwelling elephants in Asia seem to have a reduced impact on tree saplings compared to African forest elephants, but a very strong impact on monocots.     

Estimating elephant density using motion-sensitive cameras: challenges,opportunities, and parameters for consideration

With extinction rates far exceeding the natural background rate, reliable monitoring of wildlife populations has become crucial for adaptive management and conservation. Robust monitoring is often labor intensive with high economic costs, particularly in the case of those species that are subject to illegal poaching, such as elephants, which require frequent and accurate population estimates over large spatial scales. Dung counting methods are commonly employed to estimate the density of elephants; however, in the absence of a full survey calibration, these can be unreliable in heterogeneous habitats where dung decay rates may be highly variable. We explored whether motion-sensitive cameras offer a simple, lower cost, and reliable alternative for monitoring in challenging forest environments. We estimated the density of African savanna elephants (Loxodanta africana) in a montane forest using the random encounter model and assessed the importance of surveying parameters for future survey design. We deployed motion-sensitive cameras in 65 locations in the Aberdare Conservation Area in Kenya during June to August in 2015 to 2017, for a survey effort of 967 days, and a mean encounter rate of 0.09 ± 0.29 (SD) images/day. Elephants were captured in 16 locations. Density estimates varied between vegetation types, with estimates ranging from 6.27/km² in shrub, 1.1/km² in forest, 0.53/km² in bamboo (Yushania alpine), and 0.44/km² in the moorlands. The average speed of animal movement and the camera detection zone had the strongest linear associations with density estimates (R = −0.97). The random encounter model has the potential to offer an alternative, or complementary method within the active management framework for monitoring elephant populations in forests at a relatively low cost.     

The ecological basis of hunter-gatherer subsistence in African Rain Forests: The Mbuti of Eastern Zaire

The Mbuti pygmies, hunter-gatherers of the Ituri Forest of Zaire, trade forest products and labor for agricultural foods. It has been assumed that the Mbuti lived independently in the equatorial forest prior to its penetration by shifting cultivators. We assessed forest food resources (plant and animal) to determine their adequacy to support a hunting and gathering economy. For five months of the year, essentially none of the calorically important forest fruits and seeds are available. Honey is not abundant during this season of scarcity. Wild game meat is available year round, but the main animals caught have low fat content. This makes them a poor substitute for starch-dense agricultural foods, now staples in Mbuti diet. In general, in the closed evergreen forest zone, edible wild plant species are more abundant in agriculturally derived secondary forest than in primary forest. Similarly, they are more common at the savanna ecotone and in gallery forests. We suggest that it is unlikely that hunter-gatherers would have lived independently in the forest interior with its precarious resource base, when many of the food species they exploit are more abundant toward the savanna border.     

Employing participatory surveys to monitor the illegal killing of elephants across diverse land uses in Laikipia–Samburu,Kenya

Levels and trends of illegal killing of elephants are measured by the Convention on International Trade in Endangered Species (CITES) Monitoring the Illegal Killing of Elephants (MIKE) programme in sites across Africa and Asia. In the mostly unprotected Laikipia–Samburu MIKE site in northern Kenya, elephant mortality data were collected using both standard law enforcement monitoring procedures, relying on patrolling, and participatory methods involving local communities. Qualitatively, traditional patrolling techniques were more successful in protected areas whereas participatory approaches provided more information outside protected areas, where elephant were most at risk from ivory poachers. A minimum of 35% of the 389 verified carcasses during 2001–2003 were illegally killed. In this baseline study, land uses ranked from highest to lowest by the proportion of illegally killed elephants (PIKE) were community conservation areas, government trust lands, forest reserves, private ranches, settlement areas and national reserves. PIKE trends derived from traditional and participatory data sources were similar across years and indicate elephants were at greater risk in regions outside government or privately patrolled areas. We suggest that PIKE is a useful index for comparing levels and trends in illegal killing of elephants, and that carcass ratios and presence/absence of tusks are useful proxy indicators of mortality and its causes.     

Megafaunal influences on tree recruitment in African equatorial forests

The forests of central Africa are distinct from counterpart forests in Amazonia by having fewer trees ≥ 10 cm dbh ha^?1, especially small trees < 20 cm dbh, and in having sapling cohorts with less diversity than canopy trees. We tested four hypotheses to investigate whether herbivory, in particular, browsing by forest elephants, could be a factor in these differences. We found that the density of small saplings and diversity of large saplings were inversely associated with local density of elephants. We then tested the hypothesis that steep slopes might serve as refugia from elephant foraging, but found that elephants routinely forage on slopes with an inclination of less than ca 30 deg. Nevertheless, the diversity of small trees (≥ 10 cm, < 20 cm dbh) was higher on slopes than on paired level‐ground sites. The incidence of break scars on saplings ≥ 2 cm dbh and < 6 cm dbh was greater (107/100 stems) on level ground than on slopes (77/100 stems), although high variability precluded statistical significance. After correcting for background breakage not caused by elephants, an estimated 71% of breaks on level ground and 43% of breaks on slopes were attributable to elephants. Liana loads borne by trees at different sites were highly variable and unrelated to slope. Apparently, disturbances are more critical to liana development than herbivory. Elephants, along with other large mammals such as gorillas, duikers, red river hogs and rodents, appear to act as powerful filters on the tree recruitment process in African forests that still retain intact megafaunal communities.     

Levels of aloe mortality with and without elephants in the Thicket Biome of South Africa

Studies concerning the influence of African elephants ( Loxodonta africana ) on vegetation have produced contradictory results; some show minimal or no effect while others report significant elephant-induced effects. Elephants are generalist megaherbivores but will selectively feed from preferred plant species. We investigated the mortality of aloe plants (highly preferred food items for elephants) at five sites with elephants (treatment) and five paired sites without elephants (control) in the Eastern Cape Province of South Africa. A significantly higher proportion of aloes were dead at treatment sites and significantly more aloes that had lost their crown (headless) were found at treatment sites compared with controls. We conclude that although the proportions of dead aloes at treatment sites were significantly higher, it remains unclear whether there is a need to be concerned with the potential small-scale extinction of aloes from parts of the Eastern Cape Province. The observed mortality may merely be an artefact of the loss of large herbivores through disease (e.g. rinderpest) and hunting in the past.     

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.     

Forest elephant crisis in the Congo Basin

Debate over repealing the ivory trade ban dominates conferences of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). Resolving this controversy requires accurate estimates of elephant population trends and rates of illegal killing. Most African savannah elephant populations are well known; however, the status of forest elephants, perhaps a distinct species, in the vast Congo Basin is unclear. We assessed population status and incidence of poaching from line-transect and reconnaissance surveys conducted on foot in sites throughout the Congo Basin. Results indicate that the abundance and range of forest elephants are threatened from poaching that is most intense close to roads. The probability of elephant presence increased with distance to roads, whereas that of human signs declined. At all distances from roads, the probability of elephant occurrence was always higher inside, compared to outside, protected areas, whereas that of humans was always lower. Inside protected areas, forest elephant density was correlated with the size of remote forest core, but not with size of protected area. Forest elephants must be prioritised in elephant management planning at the continental scale.     

Genetic connectivity across marginal habitats: the elephants of the Namib Desert

Locally isolated populations in marginal habitats may be genetically distinctive and of heightened conservation concern. Elephants inhabiting the Namib Desert have been reported to show distinctive behavioral and phenotypic adaptations in that severely arid environment. The genetic distinctiveness of Namibian desert elephants relative to other African savanna elephant (Loxodonta africana) populations has not been established. To investigate the genetic structure of elephants in Namibia, we determined the mitochondrial (mt) DNA control region sequences and genotyped 17 microsatellite loci in desert elephants (n = 8) from the Hoanib River catchment and the Hoarusib River catchment. We compared these to the genotypes of elephants (n = 77) from other localities in Namibia. The mtDNA haplotype sequences and frequencies among desert elephants were similar to those of elephants in Etosha National Park, the Huab River catchment, the Ugab River catchment, and central Kunene, although the geographically distant Caprivi Strip had different mtDNA haplotypes. Likewise, analysis of the microsatellite genotypes of desert‐dwelling elephants revealed that they were not genetically distinctive from Etosha elephants, and there was no evidence for isolation by distance across the Etosha region. These results, and a review of the historical record, suggest that a high learning capacity and long‐distance migrations allowed Namibian elephants to regularly shift their ranges to survive in the face of high variability in climate and in hunting pressure.     

Survey of Loxodonta africana (Elephantidae)‐caused bark injury on Adansonia digitata (Malavaceae) within Pendjari Biosphere Reserve,Benin

This study assessed the level of bark damage on baobab trees (Adansonia digitata) as caused by elephants (Loxodonta africana), and the possibility of finding refuges where baobab could escape bark damage within the Pendjari Biosphere Reserve (PBR). Distributions of elephants and baobab trees within the PBR were compared using presence records of both species taken along transect lines. Two sites (National Park vs. hunting zone) that differ in elephant density were compared for intensity of bark damage and correlations between the intensity of bark damage and stem size of the baobab trees and population structure of the baobab trees. Elephants and baobabs showed co‐occurrence in PBR suggesting that there is nowhere to hide for baobabs. The intensity of bark damage was positively correlated with elephant density and baobab girth. Baobab population girth classes were not significantly different in areas with and without bark damage. Future studies should test whether there are certain baobab genotypes that can resist elephant damage. It could also be tested whether effective conservation of elephants in the PBR has resulted in a bull‐biased population over its carrying capacity.     

The Human Ecology of World Systems in East Africa: The Impact of the Ivory Trade

The impact on human ecology of the ivory trade entailed direct and indirect effects. First, the reduction or extermination of elephant populations had direct effects on the vegetation patterns over large areas. Second, the economic activities connected with hunting, transport, and trading affected regional systems of exchange and thereby, indirectly through the political economy, settlements, patterns of resource utilization, population parameters, and specialization of production. Ethnohistorical information from the 1800s suggests how coastal goods interacted with regional systems of exchange and environmental exploitation. Although such information cannot be directly projected onto the more distant past, it can be used to establish some possible pathways through which the hunting of elephants and transportation and trade of ivory could have affected the ecology of human resource use.     

The Asian elephants-driver partnership: the drivers' perspective

Elephant drivers, sometimes termed mahouts, are known to share a relationship with their elephants rarely matched in other human-animal interactions with regard to time invested, extent of cooperative activity, and everpresent risk to the driver. An investigation of this relationship was pursued at two tourist lodges in Nepal where elephants are used to transport tourists into a nearby jungle to view wildlife. The study sought to investigate the drivers' perceptions regarding the individual and social behavior of the elephants, the perceptions of the elephants, and the elephants' interactions with drivers. Standardized open-ended questions were administered with translator assistance to 17 head drivers of elephants. Drivers attributed their management success to the time and care they invested in caring for and becoming familiar with the elephant. Drivers worked in partnership with elephants to gather and prepare the elephants' food. Elephants responded to vocal commands of drivers for saddling. Drivers also took responsibility for elephants in their varied interactions with tourists. Although drivers varied in specifying the most desirable elephant at their lodge, they highly agreed on the identity of the worst elephant because of its aggressivity. In general, drivers believed that their elephants loved and trusted them. Most drivers reported that their elephants did not get angry with them. Yet, they knew that elephants would most like to be free in the jungle. Drivers presented consistent information as to the elephants' social preferences for and dislikes of one another.     

African elephants show high levels of interest in the skulls and ivory of their own species

An important area of biology involves investigating the origins in animals of traits that are thought of as uniquely human. One way that humans appear unique is in the importance they attach to the dead bodies of other humans, particularly those of their close kin, and the rituals that they have developed for burying them. In contrast, most animals appear to show only limited interest in the carcasses or associated remains of dead individuals of their own species. African elephants (Loxodonta africana) are unusual in that they not only give dramatic reactions to the dead bodies of other elephants, but are also reported to systematically investigate elephant bones and tusks that they encounter, and it has sometimes been suggested that they visit the bones of relatives. Here, we use systematic presentations of object arrays to demonstrate that African elephants show higher levels of interest in elephant skulls and ivory than in natural objects or the skulls of other large terrestrial mammals. However, they do not appear to specifically select the skulls of their own relatives for investigation so that visits to dead relatives probably result from a more general attraction to elephant remains.     

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.