New evidence for hybrid zones of forest and savanna elephants in Central and West Africa

The African elephant consists of forest and savanna subspecies. Both subspecies are highly endangered due to severe poaching and habitat loss, and knowledge of their population structure is vital to their conservation. Previous studies have demonstrated marked genetic and morphological differences between forest and savanna elephants, and despite extensive sampling, genetic evidence of hybridization between them has been restricted largely to a few hybrids in the Garamba region of northeastern Democratic Republic of Congo (DRC). Here, we present new genetic data on hybridization from previously unsampled areas of Africa. Novel statistical methods applied to these data identify 46 hybrid samples – many more than have been previously identified – only two of which are from the Garamba region. The remaining 44 are from three other geographically distinct locations: a major hybrid zone along the border of the DRC and Uganda, a second potential hybrid zone in Central African Republic and a smaller fraction of hybrids in the Pendjari–Arli complex of West Africa. Most of the hybrids show evidence of interbreeding over more than one generation, demonstrating that hybrids are fertile. Mitochondrial and Y chromosome data demonstrate that the hybridization is bidirectional, involving males and females from both subspecies. We hypothesize that the hybrid zones may have been facilitated by poaching and habitat modification. The localized geography and rarity of hybrid zones, their possible facilitation from human pressures, and the high divergence and genetic distinctness of forest and savanna elephants throughout their ranges, are consistent with calls for separate species classification.     

Structure of population,grouping patterns and density of forest elephants in north-west Congo

The structure of the population of forest elephants visiting a clearing in north-west Congo was studied over 8 months. Out of 3314 sightings, 629 elephants were identified (including 64%, of adults). The sex ratio was about 1:1. The number of offspring per female was 1.12. Solitary elephants (91.4%, of males) made up 35% of the population. Of females, 93.6% were grouped. Groups (mean size 3.5) included females and offspring (3.4), males and females (5.0), or males only (2.3). The return rate of elephants at the clearing suggested that up to 1900 individuals have visited the clearing. This confirms that elephant densities are especially high in north Congo.     

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.     

Movement patterns of forest elephants (Loxodonta cyclotis Matschie, 1900) in the Odzala-Kokoua National Park,Republic of Congo

African forest elephants (Loxodonta cyclotis Matschie, 1900) are ecological engineers that play a fundamental role in vegetation dynamics. The species is of immediate conservation concern, yet it is relatively understudied. To narrow this knowledge gap, we studied the drivers of daily movement patterns (linear displacements) of forest elephants—characterised by a set of geographical, meteorological and anthropogenic variables—in the Odzala-Kokoua National Park, Republic of Congo. Explicitly, we used conditional random forest to model and disentangle the main environmental factors governing the displacements of six forest elephants, fitted with GPS collars and tracked over 16 months. Results indicated that females moved further distances than males, while the presence of roads or human settlements disrupted elephant behaviour resulting in faster displacements. Forest elephants moved faster along watercourses and through forest with understory dominated by Marantaceae forests and bais, but moved slower in savannahs. Finally, flood-prone areas—described by elevation and accumulated precipitation—and higher temperatures prevented longer displacements. We expect these results to improve the knowledge on the species movements through different habitats, which would benefit its conservation management.     

Counting elephants in Montane forests: some sources of error

The dung count method is widely used to estimate elephant numbers in forests. It was developed in the lowland forests of Central Africa but it is also used in Montane forests in eastern Africa. Using data collected on Mount Kenya and computer simulations, this paper explores the following issues associated with dung surveys in Montane forests:

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  High rainfall at 3000 m altitude on Mount Kenya was expected to accelerate dung pile decay but no significant difference was found between 3000 and 2500 m where less rain falls, possibly because high rainfall at 3000 m is counteracted by lower temperatures;
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  Physical obstacles make it difficult to walk long, straight transects in Montane forests. Deviating from a straight line pushes the distribution of distance measurements from dung piles to the transect centre line (pdist) towards a negative exponential (NE), which complicates data analysis and may give inaccurate estimates. Using short transects largely alleviate this problem;
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  Analysis of dung count simulations shows that the expected sightability curve of pdist pushes towards a NE with increasing numbers of obstacles blocking the view, even along perfectly straight transects;
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  Extrapolating measured dung density to map area on Mount Kenya resulted in an underestimate of c. 13%. An unstratified correction of map area to ground area for Montane areas would be biased because of the strong tendency for elephants to avoid steeply sloping areas.

     

Refugia, dispersal and divergence in a forest archipelago: a study of Streptocarpus in eastern South Africa

We describe a scenario of plant speciation across a relict forest archipelago in South Africa involving Pleistocene habitat expansion-contraction cycles, dispersal and adaptation to lower temperatures. This is the first population level study using molecular data in South African forests and has significant implications for conservation efforts in this area. Populations of the mesophytic forest floor herbs Streptocarpus primulifolius sensu lato and Streptocarpus rexii were sampled throughout their range in the naturally fragmented forests of eastern South Africa in order to investigate population genetic and phylogenetic patterns within the species complex, using nuclear microsatellites, nuclear ribosomal ITS (internal transcribed spacer) sequences and chloroplast genome sequences. S. primulifolius harbours high levels of genetic diversity at both the nuclear (mean HE = 0.50) and the chloroplast level (each population fixed for a unique haplotype). This is consistent with populations of these coastal species being Pleistocene relicts. In contrast, populations of S. rexii in cooler habitats at higher altitudes and lower latitudes harbour little or no nuclear genetic diversity (mean HE = 0.09) and most share a common chloroplast haplotype. The split of S. rexii from populations intermediate between the two species (S. cf. primulifolius) occurred between 0 and 0.44 million years ago according to the calibrated ITS phylogeny of the taxa. The low genetic diversity and homogeneity of S. rexii is congruent with this species having reached its current range during the Holocene. We found no evidence of monophyly of any of the taxa in this study, which we consider a consequence of recent evolution in a fragmented habitat.     

Human impact on wildlife populations within a protected Central African forest

This paper addresses the effect of human activities on the density of large mammals in the Dzanga‐Ndoki National Park and the adjacent Dzanga‐Sangha Reserve in the Central African Republic. Between six and eight 20 km long permanent transects were walked on a monthly basis from January 1997 to August 1999 to assess large mammal populations as well as human intrusion. There were no obvious seasonal or monthly trends in elephant, gorilla or non‐human primate densities. Overall, it appears that human activities negatively influence the distribution of most of the large forest animals in Dzanga‐Sangha. Elephants in particular were significantly less common in areas used by humans, but also other species such as non‐human primates showed lower densities closer to the main road and the town of Bayanga. This study confirms the findings of previous studies that roads have a negative impact on wildlife populations. Results of this study stress the need for conservation of large uninterrupted forest blocks to maintain wildlife populations at normal levels. Simply creating roads, even within a protected Central African forest, is likely to have negative impacts on wildlife populations.     

Priority areas for the conservation of subtropical indigenous forest in southern Africa: a case study from KwaZulu-Natal

Southern Africa's subtropical forest biome, though small and highly fragmented, supports much of the region's biodiversity. With limited resources available for conservation and the exploitative use of forest escalating, identifying a network of priority forest reserves is important. We examine the distribution of forest birds, butterflies and mammals in KwaZulu-Natal, South Africa. Using an iterative algorithm we explore the efficiency of existing protected areas, species richness and rarity hotspots, prime forest sites (selected by forest area) and complementary networks as alternative approaches to priority reserve selection, as well as the potential use of indicator taxa. Existing protected areas represent 98% of species but are relatively inefficient in terms of area. Alternative selection criteria represent a high proportion of species (86–92%) and provide efficient bases for developing fully representative reserve networks. All species are represented within a network of 22 complementary quarter degree cells. This network includes several larger forests and existing protected areas and is recommended for priority conservation. Complementary networks identified separately for birds, butterflies and mammals overlap little, but each represents a high proportion of the remaining taxa, supporting their potential as representative 'indicator' taxa. The evolutionary history of the three main forest types in KwaZulu-Natal explains observed spatial patterns of alternative reserve networks. Priority areas are concentrated in scarp and coastal forest belts, regions of comparatively recent evolutionary activity with high species richness. Afromontane forest is older and less diverse, but its inclusion in any reserve network is necessary for the full representation of forest diversity.     

Man determines the distribution of elephants in the rain forests of northeastern Gabon

Dropping counts were used to assess elephant abundance in the remote forests of northeastern Gabon where there are few people and no logging. Elephants prefer the secondary forest which grows on abandoned villages and plantations, but avoid roads and villages. Thus elephant distribution is governed by the distribution of both past and present human settlement, even in the remotest and least disturbed forests of equatorial Africa.

Résumé

On a évalué l'abondance des éléphants en comptant les excréments, dans les forêts reculées du nord-est du Gabon, là où il n'y a que peu de gens et aucun abattage d'arbres. Les éléphants préférent les forêts secondaires qui poussent dans les villages et les plantations abandonnés, mais évitent routes et villages. Donc, la distribution des éléphants est régie par la répartition des installations humaines présentes et passées, même dans les forêts les plus reculées et les moins troublées d'Afrique équatoriale.     

The ranging patterns of elephants in Marsabit protected area,Kenya: the use of satellite‐linked GPS collars

We investigated the ranging patterns of elephants in the Marsabit protected area, north eastern Kenya, to ascertain the range of bachelor and female family herds in different seasons, and to identify corridor and noncorridor areas. Data were acquired for five bachelor and four female family herds equipped with satellite‐linked geographical positioning system collars, and monitored from December 2005 to December 2007. Distinct dry (about 260 km²) and wet seasons (about 910 km²) ranges were observed, with connecting corridors (north‐eastern corridor: about 90 km long, about 2‐7 km wide; southern corridors: about 10‐20 km long, about 2‐3 km wide). The dry season range corresponded with Marsabit evergreen forest, while the wet season range matched with dry deciduous lowland shrubs. The ranging elephants moved at speed of about 0.2‐20 kmh^?1. Bachelor herds moved faster than female family herds. Elephants moved fast during the intermediate and wet seasons than during the dry season. The speed of ranging elephants was over 1 kmh^?1 in the corridor areas and about 0.2 to less than 1 kmh^?1 in the non‐corridor areas. Expansion of settlements towards corridor areas needs to be controlled to avoid future blocking of connectivity between wet and dry season elephant ranges.     

Spatial and temporal variability in the structure of a tropical forest

This study examines spatial and temporal variation in the forest structure of the Kibale National Park, Uganda by contrasting tree density, tree size, and forest composition among four areas each separated by less than 15 km, and by quantifying changes in the composition of one of these forests over a 20‐year period. Densities of some tree species differed markedly between sites, and some species common at one location were absent at others. Monthly phenological monitoring demonstrated that it was not uncommon for phenological patterns to differ between the forests. To examine temporal variation in the tree composition over a 20‐year period, a sampling regime that was carried out in the early 1970s was replicated on the floristic composition of one of these sites, using identical methods in the same sampling areas. While no form of human intervention occurred in this area between the early 1970s and 1992, there were marked changes in the densities of some tree species. Twenty‐seven percent of the identified species increased in abundance, 33% decreased, and 40% remained relatively unchanged. The observed spatial and temporal variation in forest composition could be the result of abiotic factors, such as altitude or rainfall, or biotic factors such as elephant and/or human influences on ecosystem dynamics; the implications of this variation for frugivores are discussed.     

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.     

Modeling population viability of captive elephants in Myanmar (Burma): implications for wild populations

Captive Asian elephants Elephas maximus , used as work animals, constitute up to 22–30% of remaining Asian elephants. Myanmar has the largest captive population worldwide (∼6000), maintained at this level for over a century. We used published demographic data to assess the viability of this captive population. We tested how this population can be self-sustained, how many elephants must be supplemented from the wild to maintain it, and what consequences live capture may have for Myanmar's wild population. Our results demonstrate that the current captive population is not self-sustaining because mortality is too high and birth rates are too low. Our models also suggest ∼100 elephants year⁻¹ have been captured in the wild to supplement the captive population. Such supplementation cannot be supported by a wild population of fewer than 4000 elephants. Given the most recent expert estimate of ∼2000 wild elephants remaining in Myanmar, a harvest of 100 elephants year⁻¹ could result in extinction of the wild population in 31 years. Continued live capture threatens the survival of wild and captive populations and must stop. In addition, captive breeding should be increased. These measures are essential to slow the decline and extinction of all of Myanmar's elephants.     

Birth seasonality and calf mortality in a large population of Asian elephants

In seasonal environments, many species concentrate their reproduction in the time of year most likely to maximize offspring survival. Asian elephants (Elephas maximus) inhabit regions with seasonal climate, but females can still experience 16‐week reproductive cycles throughout the year. Whether female elephants nevertheless concentrate births on periods with maximum offspring survival prospects remains unknown. We investigated the seasonal timing of births, and effects of birth month on short‐ and long‐term mortality of Asian elephants, using a unique demographic data set of 2350 semicaptive, longitudinally monitored logging elephants from Myanmar experiencing seasonal variation in both workload and environmental conditions. Our results show variation in birth rate across the year, with 41% of births occurring between December and March. This corresponds to the cool, dry period and the beginning of the hot season, and to conceptions occurring during the resting, nonlogging period between February and June. Giving birth during the peak December to March period improves offspring survival, as the odds for survival between age 1 and 5 years are 44% higher for individuals born during the high birth rate period than those conceived during working months. Our results suggest that seasonal conditions, most likely maternal workload and/or climate, limit conception rate and calf survival in this population through effects on maternal stress, estrus cycles, or access to mates. This has implications for improving the birth rate and infant survival in captive populations by limiting workload of females of reproductive age. As working populations are currently unsustainable and supplemented through the capture of wild elephants, it is imperative to the conservation of Asian elephants to understand and alleviate the effects of seasonal conditions on vital rates in the working population in order to reduce the pressure for further capture from the wild.