African Savannah Elephants Research Articles

Modeling harvest rates and numbers from age and sex ratios: A demonstration for elephant populations

Illegal harvest rates of wildlife populations are often unknown or difficult to estimate from field data due to under-reporting or incomplete detection of carcasses. This is especially true for elephants that are killed for ivory or in conflicts with people. We describe a method to infer harvest rates from coarse field data of three population parameters, namely, adult female to male ratio, male old-adult to young-adult ratio, and proportion of adult males in the population using Jensen’s (2000) 2-sex, density-dependent Leslie matrix model. The specific combination of male and female harvest rates and numbers can be determined from the history of harvest and estimate of population size. We applied this technique to two populations of elephants for which data on age structure and records of mortality were available—a forest-dwelling population of the Asian elephant (at Nagarahole, India) and an African savannah elephant population (at Samburu, Kenya) that had experienced male-biased harvest regimes over 2–3decades. For the Nagarahole population, the recorded numbers of male and female elephants killed illegally during 1981–2000 were 64% and 88% of the values predicted by the model, respectively, implying some non-detection or incomplete reporting while for the Samburu population the recorded and modeled numbers of harvest during 1990–1999 closely matched. This technique, applicable to any animal population following logistic growth model, can be especially useful for inferring illegal harvest numbers of forest elephants in Africa and Asia.

Forest elephants: fission–fusion and social arenas

Fission–fusion systems provide models for studying grouping decisions and social complexity in a range of taxa, as individuals act to manage risks and enhance reproductive success. African savannah elephants, Loxodonta africana africana , live in a fluid society where individual relationships persist over decades and affect reproductive success. Social structures have yet to be fully characterized for forest elephants, Loxodonta africana cyclotis . Aggregations at natural forest clearings, which are frequent in this subspecies, may offer social benefits in addition to nutritional resources. We examined the social functions of aggregations in Odzala–Kokoua National Park, Republic of Congo. We tested whether clearings function as ‘social arenas’ by examining how social opportunities affected the timing and duration of visits, and how short-term grouping decisions were affected by age, sex, group size and number of elephants present. Aggregations reflected a fission–fusion society similar to that observed in savannah elephants. Grouping patterns were largely unaffected by the number of elephants present, from which we infer that elephants were not acting to minimize competition. Social opportunities affected visit patterns such that visits were longer when others were present. Social opportunities were maximized by joining large groups, often including older elephants, or, for males, parties with females. Large and dynamically changing social groups in clearings offer potential for information exchange and the maintenance of social relationships. Demonstrating that fission–fusion operates in forest elephant society allows future research to focus on cost–benefit modelling, to understand the fitness consequences of these groupings.

GENETIC IDENTIFICATION OF FIVE STRONGYLE NEMATODE PARASITES IN WILD AFRICAN ELEPHANTS (LOXODONTA AFRICANA)

African savannah elephants (Loxodonta africana) are an ecologically and economically important species in many African habitats. However, despite the importance of elephants, research on their parasites is limited, especially in wild populations. Currently, we lack genetic tools to identify elephant parasites. We present genetic markers from ribosomal DNA (rDNA) and mitochondrial DNA (mtDNA) to identify five elephant-specific nematode parasites in the family Strongylidae: Murshidia linstowi, Murshidia longicaudata, Murshidia africana, Quilonia africana, and Khalilia sameera. We collected adult nematodes from feces deposited by wild elephants living in Amboseli National Park, Kenya. Using both morphologic and genetic techniques, we found that the internal transcribed spacer (ITS) region in rDNA provides a reliable marker to distinguish these species of strongyles. We found no evidence for cryptic genetic species within these morphologic species according to the cox-1 region of mtDNA. Levels of genetic diversity in strongyles from elephants were consistent with the genetic diversity seen within other strongyle species. We anticipate that these results will be a useful tool for identifying gastrointestinal nematode parasites in elephants.

Functional responses in the habitat selection of a generalist mega‐herbivore, the African savannah elephant

Resource selection function (RSF) models are commonly used to quantify species/habitat associations and predict species occurrence on the landscape. However, these models are sensitive to changes in resource availability and can result in a functional response to resource abundance, where preferences change as a function of availability. For generalist species, which utilize a wide range of habitats and resources, quantifying habitat selection is particularly challenging. Spatial and temporal changes in resource abundance can result in changes in selection preference affecting the robustness of habitat selection models. We examined selection preference across a wide range of ecological conditions for a generalist mega‐herbivore, the African savanna elephant Loxodonta africana, to quantify general patterns in selection and to illustrate the importance of functional responses in elephant habitat selection. We found a functional response in habitat selection across both space and time for tree cover, with tree cover being unimportant to habitat selection in the mesic, eastern populations during the wet season. A temporal functional response for water was also evident, with greater variability in selection during the wet season. Selection for low slopes, high tree cover, and far distance from people was consistent across populations; however, variability in selection coefficients changed as a function of the abundance of a given resource within the home range. This variability of selection coefficients could be used to improve confidence estimations for inferences drawn from habitat selection models. Quantifying functional responses in habitat selection is one way to better predict how wildlife will respond to an ever‐changing environment, and they provide promising insights into the habitat selection of generalist species.

Historical range and movements of the Elephants in Babile Elephant Sanctuary, Ethiopia

AbstractA study on historical range and movement patterns of the African savannah elephants (Loxodonta africana africana) in Ethiopia was made from December 2004 to November 2008. Babile Elephant Sanctuary is one of the strongholds of elephants in the country, where it now supports approximately 27% of the elephant population in Ethiopia. Elephant movement routes and ranges were identified based on interviews of local people and by tracking the animals in the field. Elephants of the Sanctuary were categorized into two distinctive groups or clans based on their movement patterns and their associations to each other. Three major valleys, namely Gobele, Erer and Dakata, were crossed by elephants. Elephants in the population had an estimated annual range of 3014 km2. Of this, 85.7% (2583 km2) was inside the Sanctuary and 14.3% (430 km2) outside but adjacent to the Sanctuary. Elephant use outside of the Sanctuary included three different areas: Upper Erer Valley (8 km2), Upper Gobele Valley (48 km2) and west of Gobele Valley (374 km2). The extent of range varied seasonally. During the wet season, elephant range was restricted to Erer Valley and its tributaries (1266 km2 and a density of approximately 0.25 animal km−2), or 42% smaller than the dry season range (1417 km2, 0.23 elephant km−2). Range use was smallest (332 km2) in October to November and from April to May. During the dry season, elephants were found in two distinctive groups, one using Gobele Valley and the other Erer Valley.

A Comparison of Social Organization in Asian Elephants and African Savannah Elephants

Asian and African elephant species have diverged by ca. 6 million years, but as large, generalist herbivores they occupy similar niches in their respective environments. Although the multilevel, hierarchical nature of African savannah elephant societies is well established, it has been unclear whether Asian elephants behave similarly. Here we quantitatively compare the structure of both species’ societies using association data collected using the same protocol over similar time periods. Sociality in both species demonstrates well-defined structure, but in contrast to the African elephants of Samburu the Uda Walawe Asian elephants are found in smaller groups, do not maintain coherent core groups, demonstrate markedly less social connectivity at the population level, and are socially less influenced by seasonal differences in ecological conditions. The Uda Walawe Asian elephants, however, do maintain a complex, well-networked society consisting of ≥2 differentiated types of associates we term ephemeral and long-term affiliates. These findings imply we must broaden our recognition of multilevel social organization to encompass societies that fall along a gradient of nestedness, and not merely those that exhibit hierarchical nesting. This in turn suggests that multilevel structures may be more diverse and widespread than generally thought, and that phylogenetic comparisons within species-rich clades, such as that of primates, using the methods presented can provide fresh insights into their socioecological basis.

A cross-species comparison of escape from X inactivation in Eutheria: implications for evolution of X chromosome inactivation

Sex chromosome dosage compensation in both eutherian and marsupial mammals is achieved by X chromosome inactivation (XCI)—transcriptional repression that silences one of the two X chromosomes in the somatic cells of females. We recently used RNA fluorescent in situ hybridization (FISH) to show, in individual nuclei, that marsupial X inactivation (in the absence of XIST) occurs on a gene-by-gene basis, and that escape from inactivation is stochastic and independent of gene location. In the absence of similar data from fibroblast cell lines of eutherian representatives, a meaningful comparison is lacking. We therefore used RNA-FISH to examine XCI in fibroblast cell lines obtained from three distantly related eutherian model species: African savannah elephant (Loxodonta africana), mouse (Mus musculus) and human (Homo sapiens). We show that, unlike the orthologous marsupial X, inactivation of the X conserved region (XCR) in eutherians generally is complete. Two-colour RNA-FISH on female human, mouse and elephant interphase nuclei showed that XCR loci have monoallelic expression in almost all nuclei. However, we found that many loci located in the evolutionarily distinct recently added region (XAR) displayed reproducible locus-specific frequencies of nuclei with either one or two active X alleles. We propose that marsupial XCI retains features of an ancient incomplete silencing mechanism that was augmented by the evolution of the XIST gene that progressively stabilized the eutherian XCR. In contrast, the recently added region of the eutherian X displays an incomplete inactivation profile similar to that observed on the evolutionarily distinct marsupial X and the independently evolved monotreme X chromosomes.Electronic supplementary materialThe online version of this article (doi:10.1007/s00412-011-0343-8) contains supplementary material, which is available to authorized users.

Megagardeners of the forest – the role of elephants in seed dispersal

As the largest frugivores on Earth, elephants have unique ecological properties. Found in deserts, savannahs, and forests, they are the last remnants of a diverse lineage. Among the three currently recognized forms, African forest elephants are the most frugivorous, followed by Asian and African savannah elephants, although their role as seed dispersers is variable and context-dependent. African forest elephants may consume more seeds from more species than any other taxon of large vertebrate dispersers, defecating them over long distances in viable conditions into nutrient-rich and protective dung. In short, elephants are forest gardeners. The signature of elephant dispersal is evident in the spatial distribution of trees suggesting that elephants maintain tree diversity and retain low redundancy in seed dispersal systems. Large numbers of forest elephants ranging over large areas may be essential for ecosystem function. The loss of elephants will have important negative consequences for the ecological trajectories of some plant species and whole ecological communities, yet the conservation status of forest elephants is catastrophic in Asia and rapidly becoming so in Africa due to hunting and other conflicts with people. In this paper we review the current knowledge of elephants as seed dispersers, discuss the ecological consequences of their decline, and suggest priority areas for research and conservation action.

Fences and artificial water affect African savannah elephant movement patterns

The IUCN Redlist considers the African savannah elephant ( Loxodonta africana) to be “vulnerable” despite it numbering in the 100,000s and having a large geographical range. This seeming paradox stems from how quickly human persecution can eliminate elephants across large areas and how quickly elephant numbers can increase when protected. Much elephant research concentrates on the extent and consequences of elephant persecution. Where elephants thrive, two other human interventions, the provision of artificial water and the construction of fences may have large, and perhaps unintended, impacts on elephant behavior. In general, successful management requires that we understand elephant movements and land-use choices across large areas and long periods. Here, we ask specifically how artificial water and fences might affect these movements. To do this, we first characterize how elephants move in different seasons and landscape types, in different years, and how these patterns change over a region that varies considerably in annual rainfall. We fitted 73 elephants with GPS collars across a large rainfall gradient spanning seven southern African countries over a period of 6 years. We analyzed remotely-sensed environmental data from four satellite borne sensors that measure daily rainfall, weekly temperature, bi-monthly greenness, and summarise human infrastructure. Elephants move approximately 6 km/day in dry landscapes, down to approximately 3 km/day in the wettest ones. Strong seasonal differences modulate geographic differences. Elephants move less, cover less area, and are more faithful to landscapes across years in the dry season than the wet. Water availability drives these seasonal patterns. Seasonal differences in the area covered are less pronounced in wet landscapes where permanent water is more dispersed. Within-day movements reveal that elephants are consistently crepuscular but more active at night than midday when temperatures are high. Direction-changes are centered at midnight when elephants are close to water indicating regular nighttime treks to water. By design, our analyses seek to find general patterns of elephant movements — something that one can achieve only across a large range of locations and ecological conditions — in order to understand the impact of human interventions. We show that both interventions reduce seasonal differences in elephant ranging patterns and increase local impacts of elephants on the vegetation. Artificial water sources allow more extensive dry season ranging, allowing elephants to use — and potentially overexploit — vegetation in areas that would have be otherwise inaccessible to them except in the wet season. Fences cause elephants to “bunch-up” against them during the wet season, again locally increasing the pressure elephants put on their resources.

Ivory identification by DNA profiling of cytochrome b gene

Ivory can be visually identified in its native form as coming from an elephant species; however, determining from which of the three extant elephant species a section of ivory originates is more problematic. We report on a method that will identify and distinguish the protected and endangered elephant species, Elephas maximus or Loxodonta sp. To identify the species of elephant from ivory products, we developed three groups of nested PCR amplifications within the cytochrome b gene that generate amplification products using highly degraded DNA isolated from confiscated ivory samples dating from 1995. DNA from a total of 382 out of 453 ivory samples were successfully isolated and amplified leading to species identification. All sequences were searched against GenBank and found to match with E. maximus and Loxodonta sp. with at least 99% similarity. The samples that were tested came from eight Asian elephants, 14 African forest elephants (Loxodonta cyclotis), and 360 African savannah elephants (Loxodonta africana). This study demonstrates a high success rate in species identification of ivory by a nested PCR approach within the cytochrome b gene which provides the necessary information for the protection of endangered species conservation.

Elephant cognition

Elephant cognition

Population Genetic Structure of Savannah Elephants in Kenya: Conservation and Management Implications

We investigated population genetic structure and regional differentiation among African savannah elephants in Kenya using mitochondrial and microsatellite markers. We observed mitochondrial DNA (mtDNA) nucleotide diversity of 1.68% and microsatellite variation in terms of average number of alleles, expected and observed heterozygosities in the total study population of 10.20, 0.75, and 0.69, respectively. Hierarchical analysis of molecular variance of mtDNA variation revealed significant differentiation among the 3 geographical regions studied (F(CT) = 0.264; P < 0.05) and a relatively lower differentiation among populations within regions (F(SC) = 0.218; P < 0.0001). Microsatellite variation significantly differentiated among populations within regions (F(SC) = 0.019; P < 0.0001) but not at the regional levels (F(CT) = 0.000; P > 0.500). We attribute the high differentiation at the mitochondrial genome to the matrilineal social structure of elephant populations, female natal philopatry, and probably ancient vicariance. Lack of significant regional differentiation at the nuclear loci vis-a-vis strong differences at mtDNA loci between regions is likely the effect of subsequent homogenization through male-mediated gene flow. Our results depicting 3 broad regional mtDNA groups and the observed population genetic differentiation as well as connectivity patterns should be incorporated in the planning of future management activities such as translocations.

A Rapid Method to Estimate Population Variables for African Elephants

Abstract: We developed a noninvasive method to estimate reproductive and survival parameters for free‐ranging African savannah elephants (Loxodonta africana africana) and used these to estimate finite population growth rates. We used published data from 2 populations with known growth rates and birth and survival histories to validate our technique. Based on body measurements, our method yielded estimates of age at first and last calving, calving interval, and age‐specific survival rates that were similar to those determined during long‐term studies at both Addo Elephant National Park and Amboseli National Park. Our technique generated population data required to estimate population growth rates. The method may be particularly useful where censuses yield imprecise estimates or where long‐term population data are unavailable. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):822–829; 2008)

Call repertoire of infant African elephants: First insights into the early vocal ontogeny

African savannah elephants (Loxodonta africana) have a complex acoustic communication system, but very little is known about their vocal ontogeny. A first approach in ontogenetic studies is to define the call repertoire of specific age groups. Twelve hundred calls of 11 infant elephants from neonatal to 18 months of age recorded at the Vienna Zoo in Austria and at the Daphne Sheldrick's orphanage at the Nairobi National Park, Kenya were analyzed. Six call types were structurally distinguished: the rumble, the bark, the grunt, the roar (subdivided into a noisy-, tonal-, and mixed-roar), the snort, and the trumpet. Generally, within-call-type variation was high in all individuals. In contrast to adult elephants, the infants showed no gender-dependent variation in the structure or in the number of call types produced. Male infants, however, were more vocally adamant in their suckle behavior than females. These results give a first insight to the early vocal ontogeny and should promote further ontogenetic studies on elephants. Due to their vocal learning ability in combination with the complex fission-fusion society, elephants could be an interesting model to study the role of imitation in the vocal ontogeny of a nonprimate terrestrial mammal.

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.

The African elephant – something to cherish and to use

The African savannah (bush) elephant Loxodonta africana is a wonderful animal, yet at the same time it is a problem both to the environment and to rural people. Aspects of its biology are listed, and its valuable products are noted. These latter are its ecotourism potential, its trophy (some spectacular tusks are tabulated), the ivory itself, skin, curios, and meat. The ‘elephant problem’ became apparent in Africa in the early 1960s: the bulls in particular can severely damage trees, and this effect is seen from an elephant density of 0.5 per sq. km. Throughout Africa, with the exception of southern Africa, poaching has been rampant and has culled the populations. Management aspects, such as legal culling, are considered. Ecotourism dollars should be re‐invested in the resource, and sustainable use of the animal should be the governing idea.

The African savannah elephant population in Kidepo Valley National Park, Uganda: changes in size and structure from 1967 to 2000

AbstractThe age and sex structure of the elephant population in Kidepo Valley National Park were studied using recognition files. Moreover, population trends were reviewed using past studies. From 1967 to 2000, the elephant population varied between 200 and 500 individuals. Of the minimum number of 374 elephants found to use the Park, 352 were individually identified. Seventy‐nine per cent of these were recorded in 29 families ranging from three to 22 animals with a mean group size of ten. Young animals dominated the population (45% aged 0–9.9 years), while 11% were 10–14.9 years of age and only 18% over 25 years of age. Overall, males constituted 45% of the population, but only 32% of the individuals in the 10–14.9‐year age class were males. Individuals ≥15 years of age displayed a skewed sex ratio of 1 : 1.4 in favour of females. The extent of skew was greatest among mature animals (≥25 years of age), which had four times as many females as males. The skewed age structure in the year 2000 caused by poaching and drought, led to a significant difference between the age distribution in 1970 and 2000. Recent counts suggest that the Kidepo elephant population may be increasing.

The ties that bind: genetic relatedness predicts the fission and fusion of social groups in wild African elephants.

Many social animals live in stable groups. In contrast, African savannah elephants (Loxodonta africana) live in unusually fluid, fission-fusion societies. That is, 'core' social groups are composed of predictable sets of individuals; however, over the course of hours or days, these groups may temporarily divide and reunite, or they may fuse with other social groups to form much larger social units. Here, we test the hypothesis that genetic relatedness predicts patterns of group fission and fusion among wild, female African elephants. Our study of a single Kenyan population spans 236 individuals in 45 core social groups, genotyped at 11 microsatellite and one mitochondrial DNA (mtDNA) locus. We found that genetic relatedness predicted group fission; adult females remained with their first order maternal relatives when core groups fissioned temporarily. Relatedness also predicted temporary fusion between social groups; core groups were more likely to fuse with each other when the oldest females in each group were genetic relatives. Groups that shared mtDNA haplotypes were also significantly more likely to fuse than groups that did not share mtDNA. Our results suggest that associations between core social groups persist for decades after the original maternal kin have died. We discuss these results in the context of kin selection and its possible role in the evolution of elephant sociality.

Noninvasive Genotyping and Mendelian Analysis of Microsatellites in African Savannah Elephants

We obtained fresh dung samples from 202 (133 mother-offspring pairs) savannah elephants (Loxodonta africana) in Samburu, Kenya, and genotyped them at 20 microsatellite loci to assess genotyping success and errors. A total of 98.6% consensus genotypes was successfully obtained, with allelic dropout and false allele rates at 1.6% (n = 46) and 0.9% (n = 37) of heterozygous and total consensus genotypes, respectively, and an overall genotyping error rate of 2.5% based on repeat typing. Mendelian analysis revealed consistent inheritance in all but 38 allelic pairs from mother-offspring, giving an average mismatch error rate of 2.06%, a possible result of null alleles, mutations, genotyping errors, or inaccuracy in maternity assignment. We detected no evidence for large allele dropout, stuttering, or scoring error in the dataset and significant Hardy-Weinberg deviations at only two loci due to heterozygosity deficiency. Across loci, null allele frequencies were low (range: 0.000-0.042) and below the 0.20 threshold that would significantly bias individual-based studies. The high genotyping success and low errors observed in this study demonstrate reliability of the method employed and underscore the application of simple pedigrees in noninvasive studies. Since none of the sires were included in this study, the error rates presented are just estimates.

Elephants are capable of vocal learning

There are a few mammalian species that can modify their vocalizations in response to auditory experience--for example, some marine mammals use vocal imitation for reproductive advertisement, as birds sometimes do. Here we describe two examples of vocal imitation by African savannah elephants, Loxodonta africana, a terrestrial mammal that lives in a complex fission-fusion society. Our findings favour a role for vocal imitation that has already been proposed for primates, birds, bats and marine mammals: it is a useful form of acoustic communication that helps to maintain individual-specific bonds within changing social groupings.