The natural regulation of buffalo populations in East Africa: The food supply as a regulating factor,and competition*

Evidence is presented to show that both the quality and quantity of food available to a buffalo population falls below the minimum maintenance requirements of that population at certain times of the year. In the Serengeti grasslands there was a shortage of the only good quality component, grass leaf, in the dry season, with the result that the animals consumed an increasing proportion of poor quality food such as grass stem. By the end of the dry season the diet had dropped in quality below the minimum maintenance level. In areas such as Mt. Meru where there was a more continuous growing season, the high density of animals kept the standing crop of leaf at a low level. During the cooler dry season the growth of leaf became insufficient in quantity for the maintenance requirements of the population. These two quite different situations suggested that food shortage was a more general phenomenon in eastern Africa. Various measurements of feeding behaviour were made. Total grazing time per 24 h did not differ between seasons but ruminating time may have increased as the season became drier and could have been a response to the more fibrous food. Analysis for cycles of activity showed that there was more temporal organization during the dry season. These changes in activity cycles appeared to be related to the increase in energy expenditure produced by heat stress and sweating. Old animals with poor teeth did not compensate for the poor food supply by changing their feeding behaviour. There was a positive relationship between annual rainfall and mean crude density in different areas of eastern Africa, indicating that regulation was taking place. Since rainfall determined the amount of available food, it could have operated through the food supply. On a finer scale it was found that the extent of the preferred riverine habitats was also related to density. Thus rainfall, the extent of riverine habitat and perhaps soil moisture were three limiting factors that determined mean density and all could have taken effect through the food supply. As a result of initial selective grazing the amount of available leaf declined as the dry season progressed to the extent that by the end of the season the proportion of this component in the diet fell to a very low level. The impact of the population on its limited food supply indicated that intra-specific competition was acting as the cause of regulation. Measurements of wildebeest eating the same food in the same habitats as buffalo showed that inter-specific competition was also taking place. A small proportion (7“) of the wildebeest population could have reduced the buffalo population by approximately 18^o,‘, from its potential population size. The buffalo population in the Serengeti was regulated by adult mortality which was caused by undernutrition as a result of food shortage. This food shortage was caused by intra- and inter-specific competition. The effect of predation and disease was to hasten the response of the population to changes in the food supply. The limiting factors determining the mean level of the available food were, amongst others, rainfall, soil moisture and the extent of the preferred riverine habitat. The effect of interspecific competition could result in a complex regulation of populations through their food supply. There appears to be no foundation for hypotheses which invoke over-utilization or damage as a consequence of regulation through food.     

Food selection and competition in the East African buffalo (Syncerus caffer Sparrman)

Rumen samples collected at different times of the year from buffalo in the Serengeti, were analysed with respect to proportions of plant parts present. Buffalo were almost exclusive grazers and were capable of selecting for grass leaf. At certain times of year it was the dominant food intake component but declined to only 11% of the diet by the end of the dry season. The amount of leaf in the diet was determined by the amount of rainfall which governed the growth of the food grasses. Experimental preference tests with tame animals indicated that they were capable of selecting for different grass species. The preferences were, however, for those species with a higher leaf: stem ratio. The mechanism of selection is discussed. The function of the behavioural selection appeared to be concerned with maximizing the nutrient quality of the food requirements. It appeared that ecological separation had evolved through interspecific competition.     

Diet composition and habitat selection of eland in semi-arid shrubland

This study investigated the diet composition and habitat selection of eland in semi-arid shrubland, dominated by microphyllous and leptophyllous browse species offering low leaf: stem ratios. Browse (succulent, forb and woody species) contributed 94% to the annual diet of eland. The annual proportion of grass in eland diet was low (6%), even though palatable grass species were abundant in habitats favoured by eland. Most grass was eaten in the early wet season when grasses offered young green foliage. Woody species comprising dwarf shrubs and shrubs made up the bulk of the food eaten by eland. In each season, favoured woody species contributed substantial proportions to the diet of eland. Eland used the plateau habitats in the early wet season, but valleys and slopes in the late wet and dry seasons. Habitats favoured by eland contained high abundances of plants of woody species favoured by eland. Chemical analysis indicated that woody species favoured by eland offered lower total fibre contents than other woody species available to eland. The results of the study indicated that eland are browsers that select browse of low fibre content.     

Resource partitioning along multiple niche dimensions in differently sized African savanna grazers

Resource partitioning among mammalian savanna herbivores is thought to be predominantly driven by differences in body size. In general, large herbivore species utilize abundant low quality forage while small herbivores focus on scarcer high quality food items. However, in a natural system other factors such as digestive strategy, season and the presence of megaherbivores (body size > 1000 kg) are likely to complicate allometric predictions. Non‐ruminants are probably better able to cope with abundant low quality food than ruminants of the same size causing a non‐ruminant to act ‘larger’ than allometrically predicted. Also, the effect of alternating seasons with high and low food availability on diet choice and hence the competitive interactions between co‐occurring herbivores is still poorly understood. Lastly, how megaherbivores deviate from allometric predictions (based on smaller species) is still not well quantified. In this study we examine resource partitioning among three ruminant and three non‐ruminant grazers: impala, wildebeest, buffalo, warthog, zebra and white rhinoceros (megaherbivore) in the savanna of Hluhluwe iMfolozi Park, South Africa. We analysed habitat and diet overlap, specifically grass species (something not commonly investigated) and grass height eaten, in both the wet and the dry seasons. We found that habitat utilization differences among the species were generally small and did not vary between seasons. Diets within feeding patches overlapped during the wet season but highly diverged during the dry season. Body mass differences among species explained their dry season resource partitioning for all species except for comparisons with the megaherbivore (white rhino), while differences in digestive strategy were not related to niche overlap in either season. We conclude that savanna herbivores in this system coexist mostly through body size‐driven resource partitioning in the dry‐season, with the exception of the white rhino (megaherbivore).     

Water provision alters wildebeest adaptive habitat selection and resilience in the Central Kalahari

Wildlife populations in semi‐arid regions require unrestricted mobility along ecological gradients and across large landscapes to enable adaptive responses to seasonal variability and patchy resources. In the Kalahari region of Botswana, herbivore populations historically depended on seasonal access to the nutrient‐rich Schwelle area in the wet season and to water from the Boteti River during drought periods. Blue wildebeest Connochaetes taurinus in the Central Kalahari Game Reserve (CKGR) have lost access to these key habitats due to fences and encroachment of livestock and humans. We deployed satellite collars onto 10 female wildebeest in the CKGR to examine seasonal movements and habitat selection in relation to the environmental conditions and fragmented ecosystem. Wildebeest favoured open, short‐grass pan habitats in all seasons, probably in response to better forage quality and lower predation risk. The ability to remain in pan habitats during the dry season was a result of artificial water provision. A wildebeest herd that had no artificial water in its home range survived the dry season, whereas those wildebeest that were accustomed to water provision died when their water points failed in the dry season. Thus, water provision altered adaptive behaviour and reduced resilience of the population to the arid environment.     

Seasonal habitat selection and space use by a semi‐free range herbivore in a heterogeneous savanna landscape

Understanding factors that influence habitat selection in heterogeneous landscapes is fundamental for establishing realistic models on animal distribution to inform rangeland management. In this study, we tested whether seasonal variation in habitat selection within the home range of a large herbivore was influenced by constraints such as, distances from water and central place using semi‐free range cattle (Bos taurus) as a case study. We also tested whether shifts in space use over time were dependent on spatial scale and on the overall abundance of resources. We predicted that distance from water significantly influenced dry season habitat selection while the influence of the central place on habitat selection was season‐independent. We also predicted that shifts in space use over time were spatial scale‐dependent, and that large herbivores would include more diverse habitats in their home ranges during the dry season, when water and food resources are less abundant. Multinomial logit models were used to construct habitat selection models with distances from water and central place as habitat‐specific constraints. Results showed significant variations in habitat selection between the dry and wet season. As predicted, the effect of distance from central place was season‐independent, while the effect of water was not included in the top dry season models contrary to expectation. A diverse range of habitats were also selected during the dry season including agricultural fields. Results also indicated that shifts in space use were spatial scale dependent, with core areas being more sensitive to changes than the home range. In addition, shifts in space use responded to temporal changes in habitat composition. Overall, our results suggest that semi‐free range herbivores adopt different foraging strategies in response to spatial‐temporal changes in habitat availability.     

Habitat use and mating system of the houbara bustard (Chlamydotis undulata undulata) in a semi-desertic area of North Africa: implications for conservation

Studies of the movements and home-ranges of houbara bustards (Chlamydotis undulata undulata) showed sexual and seasonal differences in the use of space, with a polygynous mating system similar to an ‘exploded-lek’ or a ‘resource-defence-polygyny’, that remains undefined. We used the arthropod biomass as an index of the trophic quality of six defined habitats and we radio-tracked 7 females and 13 males to test whether sexual and seasonal variations in habitat use were related to resource availability, and to verify if critical resources for breeding females were monopolised by males. We analysed habitat selection in both sexes separately. We used the habitat type composition of buffer zones around radio-locations to study annual and seasonal habitat selection and to identify preferred habitats, using the chi-square goodness-of-fit test. Habitat use between sexes and between seasons were compared using MANOVA based on log-ratios of habitat proportions. During the year, and in each season, both sexes appeared to be significantly selective for habitats in comparison to their availability. But males avoided esparto grass, while females used all habitats. Habitat use differed between sexes in the breeding season, but not in the non-breeding season. In spring, when food resources were abundant and uniformly distributed in space, males preferred ‘temporarily flooded areas’ and females preferred ‘reg with tall perennials’ that offered both food and cover for brooding. Critical resources were not monopolised by males and the mating system fulfilled the definition of the ‘exploded-lek’. Leks are key sites for reproduction and should be considered as priority areas in further conservation plans.     

The influence of behaviour and season on habitat selection by a small mammal

This study examined the selection of habitat by Ningaui yvonneae , a small, nocturnal insectivorous marsupial of semi-arid regions of southern Australia. In addition to idengifying habitat preferences, the study was concerned with idengifying how habitats were used, and for what purpose they were valued. Triodia irritans (hummock grass) was the most preferred habitat component overall. However, other habitat components were found to be of equal or greater preference during certain seasons and for certain behaviours. The importance of considering behaviour in studies of habitat selection was discussed. Habitat selection by N. yvonneae appeared to be influenced by predation risk and energetic reward. The selection for Triodia is thought to be due to its provision of protection and foraging opportunities. However, leaf litter was preferred over Triodia for foraging, possibly because it offered greater energetic returns. To minimise predation risk while in leaf litter, ningauis remained close to Triodia and moved larger prey items to the edge of Triodia for consumption.     

The diet of Impala (Aepyceros melampus) in the Sengwa Wildlife Research Area, Rhodesia

The seasonal variation in the plant morphological part composition of the dict of Impala in the Sengwa Wildlife Research Area, Rhodesia, was determined by the analysis of rumen content samples from shot animals. The grass species composition of the diet was examined by microscopic identification of grass fragments found in the rumens. The seasonal variation in the dicotyledonous species composition of the diet was studied by direct observation of feeding animals. The protein contents of the rumen contents and faeces were used as measures of diet quality. In the wet season, grass was selected in preference to dicotyledonous plants, and grass leaf was the preferred plant part. In the late wet and early dry seasons, forbs were the principal food. The proportion of woody dicotyledons in the diet was at a maximum in the mid-dry season, when diet quality was at a minimum. Diet quality was directly related to the proportion of grass in the diet. Female Impala had a significantly higher quality diet than males, probably as a result of differential habitat selection.     

Vegetation influence on ectomycorrhizal inoculum Available to sub-arctic willow (Salix lapponum L.) planted in an upland Site

Summary

Restoration of scrub and woodland in deforested upland sites is an important conservation activity. However, little is known about the mycorrhizal colonisation potential of upland soils or the factors that influence the distribution of mycorrhizal inoculum. We investigated the effect of existing vegetation on mycorrhizal colonisation potential for a sub-arctic willow (Salix lapponum) by planting uninoculated cuttings into plotsrepresenting two upland habitats with either grassand herbs (‘grass’) or Vaccinium myrtillus (‘vaccinium’) and assessing mycorrhizal colonisation after 14 months using morphological and molecular techniques. From 40 willow cuttings (20 in each habitat), DNA sequences of rive ectomycorrhizal (EcM) fungal taxa were recovered: Laccaria proxima, Thelephora terrestris, Hebeloma sp., ‘Thelephoraceae sp.’ and ‘Pezizales sp.’. Cuttings in the ‘grass’ habitat were dominated by Laccaria proxima and ‘Pezizales sp.’ and in the ‘vaccinium’ habitat by Thelephora terrestris which was absent from the ‘grass’ habitat. There were no significant differences between habitats in frequency of EcM inoculum (overall percentage of cuttings colonised = 70%) or colonisation potential (overall mean percentage of root tips colonised per cutting = 20 %). These data suggest that the mycorrhizal colonisation potential and diversity of fungi available to willow in these upland soils are low and planted willow may benefit from inoculum enhancement.     

Sexual segregation in habitat use by elephants in Chobe National Park, Botswana

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.     

Habitat selection, diet and interspecific associations of the rufous-tailed weaver and Fischer's lovebird

We investigated habitat selection and diets of two co-occurring endemic bird species (rufous-tailed weaver, Histurgops ruficauda , Fischer's lovebird, Agapornis fischeri ) and four other species with which they associated in mixed feeding flocks (red-billed buffalo weaver, Bubalornis niger , white-headed buffalo weaver, Dinemellia dinemellii , superb starling, Lamprotornis superbus , and Hildebrandt's starling, Lamprotornis hildebrandti ) during the dry season in Serengeti National Park Tanzania. Mixed species flocks could facilitate food acquisition and/or act as anti-predator mechanisms. Five of the six species selected grassland habitat over Acacia habitat along transects. Analysis of species association, using Cole's coefficient of association, showed that both rufous-tailed weavers and superb starlings co-occurred with red-billed buffalo weavers. Superb starlings were negatively associated with Fischer's lovebird, Hildebrandt's starling and white-headed buffalo weavers. Diet analysis revealed that the rufous-tailed weaver, white-headed buffalo weaver, and red-billed buffalo weaver were generalists eating both insects and seeds, whereas Fischer's lovebird and superb starling were specialists, selecting only seeds and insects respectively. These data offer some support for the hypothesis that mixed species flocks facilitate mutual food searching.     

食物和季节因素对杂色山雀贮食行为的影响

鸟类的贮食行为受很多因素的影响,其中食物和季节是两个十分关键的因子。采用人工投食的方式研究杂色山雀冬、春季节对松子和葵花籽两种食物的贮食选择,以及食物和季节因素对贮食位点的空间分布和微生境选择的影响。结果发现,杂色山雀优先贮食松子,仅在春季贮食少量葵花籽;主要选择树皮裂缝、灌木根部、草丛、石缝和苔藓下面五种生境进行贮食;其贮食位点的空间分布呈分散状态,集中于投食点100m范围内,密度分布随食物搬运距离的增加而递减。其贮食模式和贮食位点的微生境选择均受季节因素的影响,其中,贮食模式的季节变化可能是受生境中松子这种重要食物可获得性的下降所致,而杂色山雀冬季对树皮裂缝的利用率明显高于春季,可能是与冬季的积雪覆盖限制了其对地面贮食点的利用有关。杂色山雀的就近贮食模式可能是为了增加贮食效率,关于不同个体之间贮食位点的差异,以及季节变化对贮食位点的空间分布格局的影响还需进一步研究。     

Habitat preferences,seasonal abundance and diets of rodents in Alage,Southern Ethiopia

To understand habitat preferences, seasonal abundance and diets of rodents in wet and dry season surveys were conducted in Alage, Southern Ethiopia. Sherman and snap traps were used to capture rodents from the four habitats: bushland, Acacia woodland, maize and wheat farmlands. A total of 3312 trap nights, from the four trapping habitats, yielded 776 individuals that represented 11 species of rodents. The distribution of rodents varied between habitats and seasons. Wet season rodent abundance was 52.3% while in the dry season it was 47.7%. Seasonal differences in species abundance were insignificant. Bushland habitat had high wet and dry season abundances with 137 and 211 individuals, respectively. Abundance was low in maize farm (57 individuals) in the wet season and wheat farm (10 individuals) in the dry season. Stomach content composition analysis of snap‐trapped rodents from different habitats showed differences between species and across seasons. Six rodent species were recorded as pests on the farmlands in this study area. In conclusion, variation in habitat preferences and diet of rodents in different habitats and across seasons might be due to the role of ground cover and food sources.     

Feeding of tropical freshwater fishes: seasonality in resource availability and resource use

Summary Food resources in the environment and in the diets of small fish inhabiting two water bodies in a tropical savanna were studied during both wet and dry seasons. During the wet season (high water, abundant food) most fish species in both habitats fed predominantly on vegetation-dwelling invertebrates. Most fish species switched to alternative foods (algae and detritus) following the drastic decline in invertebrate food available towards the end of the dry season. In one habitat, this change in diet was accompanied by an increase in the volume of food intake. In the second habitat, only two larger species foraged intensively, while smaller species showed low food intake or almost ceased feeding. These differences may be explained by the high risk of predation for small fish in the second habitat. Dietary overlaps among fish species were high at the end of the dry season and moderate in the wet season. However, critical analysis of such factors as food abundance, the size and number of shared prey, and diet breadth showed that all significant overlaps were ecologically unimportant i.e. there was only weak competition for food.     

Why do lizards avoid weeds?

Invasion of native habitats by exotic plants often causes reductions in faunal diversity. However, there is little direct evidence of native fauna actively avoiding invaded habitat and few quantitative studies on the mechanisms underlying such avoidance. We quantified alterations made to the composition and physical structure of an Australian tropical savanna by grader grass (Themeda quadrivalvis); an understudied invasive grass that is associated with reduced faunal abundance and diversity. We found that grader grass profoundly changed the physical structure and floral composition of tropical savanna, forming dense lawn-like monocultures unlike the native savanna. Second, we investigated the habitat preferences of small ectotherms in partially invaded habitat, using a rainbow skink (Carlia schmeltzii) as a model system and discovered that they actively avoided grader grass. Finally, we experimentally tested predictions regarding mechanisms that may have driven the avoidance of grader grass. Predation rates and food availability were not likely the cause of grader grass avoidance, because experiments using models deployed in the field showed that predation rates were higher in native grass, and collection of invertebrates in both habitats indicated that prey availability was similar. However, mesocosm experiments on habitat selection in relation to vegetation structure, along with field measures of available operative environmental temperatures, suggested that small ectotherms probably avoid grader-grass-invaded savanna due to a suboptimal thermal environment and lack of appropriate habitat structural heterogeneity.     

Prey habitat selection under shared predation: tradeoffs between risk and competition?

An individual's choice of habitat should optimize amongst conflicting demands in a way that maximizes its fitness. Habitat selection by one species will often be influenced by presence and abundance of competitors that interact directly and indirectly with each other (such as through shared predators). The optimal habitat choice will thus depend on competition for resources by other species that can also modify predation risk. It may be possible to disentangle these two effects with careful analysis of density‐dependent habitat selection by a focal prey species. We tested this conjecture by calculating habitat isodars (graphs of density assuming ideal habitat selection) of chital deer living in two adjoining dry‐forest habitats in Gir National Park and Sanctuary, western India. The habitats differed only in presence (Sanctuary) and absence (National Park) of domestic prey (cattle and buffalo). Both species are preyed on by Asiatic lions. The habitat isodar revealed at low densities, that chital live in small groups and prefer habitat co‐occupied by livestock that reduce food resources, but also reduce predation risk. At higher densities, chital form larger groups and switch their preference toward risky habitat without livestock. The switch in chital habitat use is consistent with theories predicting that prey species should trade off safety in favor of food as population density increases.     

The feeding ecology of a selective browser, the giraffe (Giraffa camelopardalis tippelskirchi)

A herbivore can manipulate certain factors of its feeding behaviour in order to achieve the metabolic requirements for reproduction. These factors include the choice of habitat in which to feed, the selection criteria for choosing food items, and the time allocated to feeding or devoted to other energy-consuming activities. The manipulation of these behavioural factors by giraffe in the Serengeti National Park, Tanzania is analysed in relation to the seasonal changes in the quantity and quality of the food resource.
The pattern of habitat choice demonstrates seasonal movements by giraffe across the drainage catena in response to differences in the rates of browse production between woodland types.
Giraffe show a positive inter-specific selection for the minor food items with a low available biomass, and utilize the dominant species in approximate proportion to their availability. Plant part selection is shown for the flushing shoots with very high protein contents.
Selection criteria include a significant selection for phosphorus, and in the dry season a selection for high energy material by breeding females. With the high nutritional qualities of the diets, negative criteria such as secondary chemical compounds are important. Patterns of browse selection are correlated with inter-specific seasonal changes in plant phenology.
Giraffe devote more time to foraging as the biomass and quality of the food declines in the dry season. Energy is conserved by minimizing energy-consuming activities at the most demanding times of day.     

Using carbon isotope analysis of the diet of two introduced Australian megaherbivores to understand Pleistocene megafaunal extinctions

Aim Australia lost a diverse assemblage of large marsupial herbivores in the late Pleistocene, with suggestions that the extinctions were biased towards browsers. In modern times two bovines, the Asian water buffalo (Bubalus bubalis) and banteng (Bos javanicus), have established feral populations in the Northern Territory, Australia. Buffalo have aggressively expanded throughout the savanna landscape, yet banteng remain near their point of introduction on the Cobourg Peninsula. We hypothesized that this difference is related to feeding ecology, possibly reflecting a legacy of the Pleistocene extinctions. Location Western Arnhem Land, Northern Territory, Australia. Methods Analysing a previously published dataset of body mass and feeding ecology of extinct and extant marsupial herbivores, we evaluated whether browsers were at greater risk of extinction than grazers. We compared the carbon isotope composition and nitrogen content of banteng and buffalo dung in order to evaluate the hypotheses that the differences in invasion success are related to feeding ecology, and that seasonal variation in browse consumption is linked to changing nutritional quality of grass. Results Controlling for body mass, the Pleistocene extinctions were clearly biased towards browsers. Introduced banteng appear to be primarily browsers, with their diets comprising 40% grass in the wet season and 15% in the late dry season. Buffalo have a more variable diet, with an increasing proportion of browse from the wet (30%) to the late dry season (75%), and can therefore be described as switching from grazer to browser. The decline of grass in the diet of both species appears to reflect the decline in the nutritional value of grass through the dry season, an inference supported by the negative relationship between δ¹³C values and the nitrogen content of dung. Main conclusions Banteng and buffalo are much larger than extant native herbivores, of which browsers are restricted to isolated rocky habitats. This suggests that banteng and buffalo have filled niches made vacant following the Pleistocene extinctions. The success of buffalo appears to be related to their greater dietary breadth, which enables them to graze and browse in eucalypt savannas, whilst the browsing banteng remain tethered to a mosaic of rain forest patches. The restriction of browsers may be a long‐range consequence of habitat transformations associated with Aboriginal landscape burning.