We report on a study conducted on free‐ranging African elephants in the woodlands of northern Botswana. We compared bull groups and family units with regard to (1) their patterns of habitat use and (2) their ranging distances from perennial water sources. During the dry season, adult males frequented more habitat types than family units, whereas family units used a wider diversity of habitats than bulls during the wet season. Bulls roamed widely (>10 km) from perennial drinking water in the dry season, when family units congregated within 3.5 km of the rivers. During the wet season, when ephemeral pans were abundant, all elephant groups were found at intermediate distances (5 km) from the rivers. The spacing of elephants in the dry season is consistent with sexual segregation but we reject the hypothesis that this is an outcome of indirect competition for food, because our concurrent studies on elephant feeding ecology found no evidence for intraspecific competition. Instead, we propose that most adult male elephants space themselves to avoid conflict with musth bulls and roam widely in the dry season between discretely distributed feeding ‘hotspots’. The small proportion of males that are in musth remain close to family units to maximize mating opportunities, and family units are unable to range far from water in the dry season. This is due to (1) comparatively high rates of water turn‐over among juveniles and lactating cows and (2) the reduced mobility of neonates.
Differences in feeding patterns of the African elephant were examined by sex and age during the dry season in a dystrophic savanna‐woodland ecosystem in northern Botswana. Adult males had the least diverse diet in terms of woody plant species, but they consumed more plant parts than family units. The diameter of stems of food plants broken or bitten off was also greater for adult males than for females and subadult males. Adult males spent more time foraging on each woody plant than did females. The number of woody plant species and individuals present were higher at feeding sites of family units than at feeding sites of adult males, indicating that family units positioned themselves at feeding sites with higher species diversity than those of males. We argue that the most likely explanation for these differences is related to the pronounced sexual size dimorphism exhibited by elephants, resulting in sex differences in browsing patterns due to the allometric relationships that govern the tolerance of herbivores for variation in diet quality. From our results this Body Size Hypothesis is accepted rather than the alternative Scramble Competition Hypothesis, which predicts that adult male elephants consume lower quality browse because they are displaced from preferred browse as an outcome of scramble competition with adult females and their offspring. If the feeding patterns of adult male elephants were affected by intersexual scramble competition, we would expect adult males to browse at a higher level in the canopy than the smaller‐bodied females and their offspring. No evidence was found for this, although adult females were found to browse at a higher level in the canopy when feeding in close proximity to subadults and juveniles than when feeding alone. Sex differences in elephant browsing patterns are, we propose, of relevance to understanding and managing elephant impacts on African woodlands.
We tested six hypotheses to explain expected geographical differences in body masses of 1,771 brown bears (Ursus arctos) from northern and southern Europe (Sweden and Norway compared with Slovenia and Croatia): Bergmann's rule, the fasting endurance hypothesis, and the dietary meat hypothesis, which predicted larger bears in the north; and hypotheses stressing the role of high primary productivity, high population density, low seasonality, and length of the growing season, which predicted larger bears in the south. Although brown bear populations in North America vary greatly in body mass, we found no significant difference in body mass between the two European populations using a new analytical approach incorporating modeled age-standardized body masses in linear models, when correcting for sex and season. The greater variation in North America may be due primarily to the presence of large bears that feed on salmon (Oncorhynchus spp.), which does not occur in Europe. Asymptotic body masses were 115 +/- 9 (SE) kg in spring and 141 +/- 9 kg in autumn for southern females, 248 +/- 25 and 243 +/- 24 kg for southern males, 96 +/- 2 and 158 +/- 4 kg for northern females, and 201 +/- 4 and 273 +/- 6 kg for northern males, respectively. Northern bears gained more body mass before hibernation and lost more during hibernation than southern bears, probably because hibernation was twice as long in the north. Northern bears gained and southern bears lost mass during the spring, perhaps due to the greater availability and use of protein-rich food in spring in the north. As reproductive success in bears is correlated with adult female body mass in interpopulation comparisons, brown bears may have relatively similar reproductive rates throughout Europe, although minimum age at primiparity and litter interval are lower in the south.
The small size of many African protected areas makes adjacent rangelands potentially important in the local survival of wild animals. In order to assess the importance of pastoral areas to wild ungulates, we studied density and habitat choice of wild ungulates and cattle in Lake Mburo National Park, Uganda, the adjacent exclusively pastoral Nshara Dairy Ranch and on private land consisting of a mixture of ranching and subsistence farms. Transects, in the three land-use zones, were walked during the wet season and the data were analysed by DISTANCE sampling technique. We found significantly higher total density of wild ungulates on the dairy ranch compared with the National Park and private land. There was no significant difference in total wild animal density between the National Park and private land. Impala (Aepyceros melampus), zebra (Equus quagga), bushbuck (Tragelaphus scriptus) and waterbuck (Kobus ellipsiprymnus) had significantly higher densities on the dairy ranch compared to the National Park. Only eland (Taurotragus oryx) density was higher in the National Park compared to private land. Wild ungulates and cattle showed a high degree of habitat overlap, generally preferring open grassland. Our study shows that high densities of wild ungulates are not necessarily associated with protected areas. Pastoral areas may be important for populations of wild herbivores during the growing season despite a pronounced presence of livestock.
Northern Botswana and adjacent areas, have the world's largest population of African elephant (Loxodonta africana). However, a 100 years ago elephants were rare following excessive hunting. Simultaneously, ungulate populations were severely reduced by decease. The ecological effects of the reduction in large herbivores must have been substantial, but are little known. Today, however, ecosystem changes following the increase in elephant numbers cause considerable concern in Botswana. This was the background for the "BONIC" project, investigating the interactions between the increasing elephant population and other ecosystem components and processes. Results confirm that the ecosystem is changing following the increase in elephant and ungulate populations, and, presumably, developing towards a situation resembling that before the reduction of large herbivores. We see no ecological reasons to artificially change elephant numbers. There are, however, economic and social reasons to control elephants, and their range in northern Botswana may have to be artificially restricted.
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