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1.
The results are presented of a 2-month study of the Mara River hippopotamus population in the Serengeti National Park and the Maasai Mara Game Reserve. A method for correcting aerial counts of hippos in turbid rivers and based on herd behaviour characteristics is described. There were an estimated 1927 hippos at an average density of 16-1/km in the area censused. The population became more dispersed and the mean group size decreased from thirty to fifteen after a rise in water level. The value of comparisons between populations in various terms of expression of density, and mean group sizes, are discussed. The amount of day living space, defined as areas preferred or utilizable by hippo during the day, and productivity of the food plants are seen to be very important factors in hippo habitat utilization, and as constraints on population increase. Hippo use of the habitat and the effects of hippo on the vegetation at various distances from the river were quantified, and a cycle of vegetational changes suggested which may be associated with the effects of hippo grazing and fire in other parts of Africa. Groups were found to consist of nearly equal numbers of males and females. There was no evidence of territorial behaviour either on land or in the water. A number of observations on social interactions and defaecating behaviour are described and indicate that olfactory communication is very important in interactions between hippos. Hippo society is unstable due to climatic influences, but a dominance hierarchy including males and females exists in the groups. Males appear less firmly attached to groups than females, but a male is normally the dominant animal in a group.  相似文献   

2.
3.
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.  相似文献   

4.
Megaherbivores perform vital ecosystem engineering roles, and have their last remaining stronghold in Africa. Of Africa's remaining megaherbivores, the common hippopotamus (Hippopotamus amphibius) has received the least scientific and conservation attention, despite how influential their ecosystem engineering activities appear to be. Given the potentially crucial ecosystem engineering influence of hippos, as well as mounting conservation concerns threatening their long-term persistence, a review of the evidence for hippos being ecosystem engineers, and the effects of their engineering, is both timely and necessary. In this review, we assess, (i) aspects of hippo biology that underlie their unique ecosystem engineering potential; (ii) evaluate hippo ecological impacts in terrestrial and aquatic environments; (iii) compare the ecosystem engineering influence of hippos to other extant African megaherbivores; (iv) evaluate factors most critical to hippo conservation and ecosystem engineering; and (v) highlight future research directions and challenges that may yield new insights into the ecological role of hippos, and of megaherbivores more broadly. We find that a variety of key life-history traits determine the hippo's unique influence, including their semi-aquatic lifestyle, large body size, specialised gut anatomy, muzzle structure, small and partially webbed feet, and highly gregarious nature. On land, hippos create grazing lawns that contain distinct plant communities and alter fire spatial extent, which shapes woody plant demographics and might assist in maintaining fire-sensitive riverine vegetation. In water, hippos deposit nutrient-rich dung, stimulating aquatic food chains and altering water chemistry and quality, impacting a host of different organisms. Hippo trampling and wallowing alters geomorphological processes, widening riverbanks, creating new river channels, and forming gullies along well-utilised hippo paths. Taken together, we propose that these myriad impacts combine to make hippos Africa's most influential megaherbivore, specifically because of the high diversity and intensity of their ecological impacts compared with other megaherbivores, and because of their unique capacity to transfer nutrients across ecosystem boundaries, enriching both terrestrial and aquatic ecosystems. Nonetheless, water pollution and extraction for agriculture and industry, erratic rainfall patterns and human–hippo conflict, threaten hippo ecosystem engineering and persistence. Therefore, we encourage greater consideration of the unique role of hippos as ecosystem engineers when considering the functional importance of megafauna in African ecosystems, and increased attention to declining hippo habitat and populations, which if unchecked could change the way in which many African ecosystems function.  相似文献   

5.
Although several aspects of the digestive physiology of the hippopotamidae-non-ruminating foregut fermenters-have been described, ingesta kinetics and passage characteristics of these species are not well understood. The most outstanding feature of the hippo digestive physiology reported so far is the very long mean ingesta retention times (MRTs) measured by Foose [Foose, T., 1982. Trophic strategies of ruminant versus nonruminant ungulates. PhD dissertation, University of Chicago, Chicago.]. Since those data had been investigated with animals without water access, we intended to measure MRT in hippos which were allowed to enter water pools during the night. MRT parameters as well as dry matter (DM) digestibility were determined in four common (Hippopotamus amphibius) and four pygmy hippos (Hexaprotodon liberiensis) on two different diets each using cobalt ethylendiamintetraacetate (Co-EDTA) as a fluid, chromium (Cr)-mordanted fibre (<2 mm) as a particle and acid detergent lignin (ADL) as an internal digestibility marker. Four of the animals additionally received cerium (Ce)-mordanted fibres (2-10 mm) as particle markers. Total MRTs for fluids and particles ranged between 20-35 and 48-106 h in the common and between 13-39 and 32-107 h in the pygmy hippos. The difference between fluid and particle retention was greater than usually reported in ruminants. Excretion patterns of the markers differed from those usually observed in ruminants but resembled those reported for macropods (kangaroos), indicating a plug-flow reactor-like physiology in the hippo forestomach (FRST). This finding complements other described similarities between the macropod and the hippo forestomach. The measurements of larger particle retention profiles suggest that in the hippo, larger particles might be excreted either faster or at the same rate as smaller particles, indicating a general difference between ruminants and hippos with respect to differential particle retention. The digestive physiology of hippos is characterised by a generally low food intake, long ingesta retention times and dry matter digestibilities lower than reported in ruminants. Moderate digestibilities in spite of long retention times might be the result of the generally high average ingesta particle size in hippos. The comparatively easy management of pygmy hippos, together with the significant correlations between food intake, MRT and digestibility in the pygmy hippos of this study, recommends this species for further studies on the interplay of these parameters in herbivore digestive physiology.  相似文献   

6.
A community of small rodents was studied in the woodland savanna of Liwonde National Park, Malawi from September 1984 to June 1985. Seven species were recorded on two 11 ha grids, using a capture-mark-release technique. Three species ( Aelhomys chrysophilus, Acomys spinosissimus and Tatera leucogasler ) comprised 78% of all trapped individuals. Average population numbers fluctuated from 3 to 17 individuals/ha. The grid with the densest vegetation supported the greatest number of individuals. Highest population numbers occurred in the mid-dry season (when the grids appeared to act as refugia) and in the early wet season (due to recruitment of young animals). Areas which had been burned during the dry season did not support as many individuals as unburnt areas. Population survival rates were low, with only about 50% of individuals surviving to the following month. The number of individuals/ha in Liwonde NP was lower than in similar habitats in other parts of eastern Africa, probably because of the shorter rainfall season and the consequent strong seasonal fluctuations in food resources, reduced opportunity for reproduction, and overall low productivity of the ecosystem.  相似文献   

7.
Lilian's Lovebird Agapornis lilianae is a small, near-threatened parrot resident in mopane Colophospermum mopane woodlands. We investigated its diet and foraging behaviour in Liwonde National Park, Malawi. We expected that Lilian's Lovebirds would show little specialisation for a particular food source but generally feed on available seeds, fruits, flowers and other items as observed in other lovebirds. Lilian's Lovebirds fed on 30 different plant species. Lilian's Lovebirds were observed feeding in six habitat types in Liwonde National Park and adjacent areas during the wet season, and four in the dry season. In the wet season lovebirds (23% of observations) foraged in grassy wetland (dambo) areas the most, whilst in the dry season they foraged in grasslands with tree cover (18%) the most. In mopane woodland, foraging flock sizes differed significantly between the wet (mean = 19.8 ± 1.0 lovebirds) and dry season (mean = 33.6 ± 2.3 lovebirds). Grass seeds were lovebirds’ main food source from December to June. The nutritional analysis of preferred foods showed that grass seeds have a relatively high protein and energy content. Grass seed availability is reduced with savanna burning and so early season burning (before May–June) in areas in and outside the park is not recommended.  相似文献   

8.
In seasonal tropical regions, rainfall and/or temporary floods during the wet season generally increase the abundance and diversity of food resources to many consumers as compared to the dry season. Therefore, seasonality can affect intraspecific competition and ecological opportunity, which are two important ecological mechanisms underlying population and individual niche variations. Here, we took advantage of the strong seasonality in the Pantanal wetlands to investigate how within‐ and between‐individual diet variations relate to seasonal population niche dynamics of the tetra fish Astyanax lacustris. We quantified dietary niche using gut contents and stable isotopes. Tetras had higher gut fullness and better body condition in the wet season, suggesting that competition is more intense in the dry season. The population niche was broader in the wet season due to an increase in diet divergence between individuals, in spite of potential stronger competition in the dry season. We posit that low ecological opportunity in the dry season limits the diversifying effect of intraspecific competition, constraining population niche expansion. Our results add new insights on how seasonality affects population and individual diets, indicating that intraspecific competition and ecological opportunity interact to determine temporal niche variations in seasonal environments.  相似文献   

9.
Monitoring abundance of threatened species is important for conservation planning. Lilian’s Lovebird Agapornis lilianae is a near-threatened small parrot found in mopane Colophospermum mopane woodland. Its population has not been investigated in any part of its range. We investigated the abundance and density of the Lilian’s Lovebird in Liwonde National Park, Malawi. Both distance sampling (line and point transects) methods and total counts (waterhole and flyway counts) were applied. The point count method gave very low numbers and was discontinued after the first year. Line transects conducted during the wet season had the highest density estimates of 17 ± 4.8 lovebirds km?2 of mopane woodland. However, number of observations per transect in each year were low. Waterhole counts had the lowest density estimates (10 ± 3.5 lovebirds km?2). Flyway counts gave an intermediate estimate (13 ± 3.0 lovebirds km?2). The total population of Lilian’s Lovebirds in Liwonde National Park is therefore estimated to be about 4 000 individuals. The use of line transect counts at the end of the rainy season is recommended for continued monitoring of Lilian’s Lovebirds abundance in Liwonde National Park.  相似文献   

10.
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 18o,‘, 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.  相似文献   

11.
Resource manipulation, such as the creation and maintenance of grazing lawns, may shape the structure of herbivore communities. We tested the hypothesis that grazing lawns contribute towards the subsistence of the Kobus kob kob in a dystrophic West African savanna, where kob and Hippopotamus amphibius both occur. Comparison of the foliage of grazing lawns and ungrazed swards shows that hippo lawns are more nutritious with regard to both structure and nutrients; kob lawns are higher in nutrients only. Up to the early dry season, hippo lawns meet kob energy and protein demand, thereafter, the shortness of the sward limits intake. Kob lawns always provide sub-maintenance values. Grazing on ungrazed swards is least profitable. We suggest that grazing lawns are essential for the daily subsistence of mesoherbivores, particularly on nutrient-poor soil, and that megaherbivores facilitate their food supply, for at least part of the year.  相似文献   

12.
Megaherbivores play a critical role in the ecology of African savannas and grasslands. In addition, these systems are forecast to experience more frequent and severe droughts as a product of changes in the global climate. Thus, the continued conservation of megaherbivores and their associated ecosystems will require a better understanding of how megaherbivores respond to drought by shifting their movement, diet and social behaviour. We address this need by investigating the factors affecting changes in the abundance of common hippopotamus (Hippopotamus amphibius; hereafter: ‘hippos’) throughout the six major rivers of Kruger National Park, South Africa, during and following the severe drought of 2015/2016. Specifically, we aimed to understand the role of two environmental characteristics that have relevance to hippos and that changed in response to drought: vegetation condition and the extent of pooled surface water. In addition, we investigated the extent to which pre-drought density affected changes in hippo abundance. Although vegetation and daytime refugia both appeared to influence pre-drought hippo abundance, these factors were less important to the change in hippo abundance related to the drought. Instead, the response to drought was most strongly related to the pre-drought abundance of hippos, where river segments supporting more than 50 individuals prior to the drought in 2015 decreased by more than half on average. Furthermore, we show that the degree of aggregation decreased from 2015 to 2016 because of the drought, but then began to increase again as the rains returned in 2017. Our results suggest that in addition to the large pools that support large aggregations of hippos in typical years, additional smaller pools are likely important for accommodating this drought-induced dispersion. However, maintaining this distribution of pools will likely become more challenging as southern Africa's population and water demands increase.  相似文献   

13.
Spatial organization and monogamy in the mara Dolichotis patagonum   总被引:1,自引:0,他引:1  
Maras, Dolichotis patagonum , were observed and radio-tracked in Argentina. They travelled as monogamous pairs that bred either alone at solitary burrows or communally at settlements where up to 29 pairs shared warrens.
Members of a mara settlement grazed within 2.5 km of the communal warrens, using intensively about 1 ha per day, within drifting daily ranges of 11 ha, seasonal ranges of 98 ha and annual ranges of 193 ha.
Their home ranges drifted continuously. Consequently, the long-term movements of neighbouring pairs overlapped substantially, but at any given moment they were territorially spaced.
Monogamous, drifting territoriality is explained by the patchy dispersion of food, the need to minimize interference competition, and a cycle of grazing and fallowing in the use of food plants.
Two hypotheses explaining the adaptive significance of settlements are evaluated: one relates to resource availability (through the indirect effect of ground water and sheep dung on vegetation); and the other relates to predation (through the protective influence of human dwellings). The size of sheep flocks grazing at outstations during January provide a measure of the resource richness of patches where maras graze, and the richness of these patches in the dry season appear to limit the number of maras breeding at each settlement during the following wet season.
Maras face extremes of resource dispersion between the wet and dry seasons: in the former, sparsely dispersed grazing and interference competition favour spacing out and territoriality; in the latter, clumping of resources facilitates pairs congregating in herds around outstations and dry lagoons. Superimposed upon the ecological factors favouring spacing out during the wet season are the sociological factors that cause the maras to den communally. The resulting compromise is a social system unique among mammals.  相似文献   

14.
In order to promote hippopotamus management in the captive and ex-situ environment, especially the control of behavioural and physiological status during breeding and lactation seasons, we conducted a preliminary study on behavioural responses of a pair of hippos including both mother and infant in Hangzhou Wildlife Park, China. The study of the captive hippos for about 1-month in the lactation season was carried out during August and September, 2009. The behavioural patterns were identified by all occurrence sampling and instantaneous scanning sampling methods with 5-10 min intervals. As a result, mother-offspring conflicts and interactions did occur throughout the whole study period. Early maternal investment showed a positive trend in activity rhythms (slope = 0.0014, Z = 0.3027, P <0.001) and a negative trend (slope = -0.0066, Z = 0.8807, P<0.001) in territorial occupation of water, all of which supported our hypotheses that the mother hippo might exert less care for the infant and cut down on her own obligations in nursing. For infant self-independence, during the whole lactation season, the primary trends of activities and territorial occupation dynamics of the infant hippo were slightly different from before, judging from linear models (slope = -0.0017, Z = 0. 3309, P<0.001). However, the frequencies of activities were not stable, especially at around 12 days of age. The trends of territorial occupation (slope = -0.0071, Z= 0. 904, P<0.001) also showed negative dynamics in water body occupation by the time the infant hippo grew up. The general trend (slope = -0.005, Z = 0.06, P<0.001) of suckling dynamics was demonstrably negative, with an upwards fluctuation at period 3 (10-15th day). This also illustrated that as the infant developed, the dependency on the mother was reduced at the end of the lactation season. In addition, a sharp decline between P3 and P4 also supported the mother-offspring conflict theory. In general, time budgets of hippos in active behaviour were (31.8±2.1)% for the mother and (32.1 ±2.6)% for the infant. Spatial distributions in water within temporal limitations were (80.1 ±2.7)% for the mother and (81.8±2.7)% for the infant. Behavioural dynamics showed strong synchronous relations between maternal investment and infant independence. Our current short-term investigation proves to be a key in management and conservation of hippopotami during the lactation season.  相似文献   

15.
Burning is commonly used in savannas to stimulate grass regrowth for grazing ungulates. We recorded the relative use of burns occurring at different stages in the seasonal cycle, as well as in different regions of the landscape by two herds of sable. We also recorded behavioural measures of foraging efficiency and faecal nutrient contents as an indication of nutrient gains. Sable consistently concentrated their grazing on burned areas provided there was sufficient green regrowth during the dry season. In these circumstances they grazed for longer per feeding station, showed a slower step rate while foraging, and shorter between‐patch moves, and a higher probability of encountering acceptable food per step taken while foraging than on unburnt areas. In the year when only a burn with insufficient regrowth was available, sable continued to forage in the area that had been burned during the previous year. Faecal crude protein was substantially higher at the end of the dry season in the year when burned areas were utilized. Accordingly early dry season fires can be important in helping sable bridge the nutritional limitations posed by the dry season, provided sufficient soil moisture remains to promote adequate grass regrowth.  相似文献   

16.
Use of poison to kill wildlife is a threat to biodiversity. Lilian's lovebirds Agapornis lilianae are among the fatalities at poisoned waterholes in Liwonde National Park, Malawi. Their population in Liwonde National Park (LNP) represents about 20% of the global population. We investigated the drinking habits of Lilian's lovebird, availability of natural waterholes and occurrence of poisoning incidents in LNP. Results showed Lilian's lovebirds congregate at waterholes in the dry season with flocks ranging from 1 to 100 individuals. Significantly larger flocks were seen in the dry than the wet season. Poisoning incidents/year ranged from 1 to 8 and were highest in the dry season. Lilian's lovebirds were killed at a mean of four poisoning incidents each year between 2000 and 2012. Number of lovebirds found dead at a poisoned pool ranged from 5 to 50 individuals. Currently about 32% of the Lilian's lovebird population is threatened by poisoning. Other species were also noted. There is a need for increased efforts in preventing this lethal activity in LNP.  相似文献   

17.
The use of landscape zones and grass species by roan antelope, a species threatened with local extirpation within South Africa's Kruger National Park, were investigated. Plant‐based observations of grazing were made within a 300 ha enclosure in the roan range, where 40 roan antelope were confined at high density in the absence of other grazers. The study spanned the dry seasons of two years, one with average rainfall and one with low rainfall. We recorded changes in the extent of grazing of different grass species, height differences between grazed and ungrazed tillers and intensity of cropping per tuft. In the average year, the grazing pressure in the bottomland grassland was twice that in the upland savanna, with two tall grass species bearing the brunt of the grazing through the dry season. Two highly palatable upland grasses were also extensively grazed by the mid dry season. In the dry year, the extent of grazing in the upland exceeded that in the bottomland, and several upland grass species little used the previous year became heavily grazed. Roan antelope appeared to be separated ecologically from more common grazers by their selective use of tall grasses growing in the drainage line grassland during the critical dry season months. However, their grazing expansion into the upland savanna during the dry year potentially brought them into competition with these grazers. Nevertheless, their population performance did not suffer despite the high‐density conditions. Heightened predation pressure following an influx of these grazers, rather than resource limitation, appeared to be primarily responsible for the drastic decline of this species in the park.  相似文献   

18.
Knowledge of a species’ movement behaviour and habitat choice is a prerequisite for assessing its ecological requirements to plan successful conservation strategies. Little is known about these factors in the Rock Firefinch, a recently described species which is probably endemic to the Jos Plateau in central Nigeria. We investigated home range size, habitat use and breeding behaviour of the Rock Firefinch in Amurum forest reserve in central Nigeria during the wet (August–October) and the dry season (November–December) using radiotracking. Birds showed high site fidelity. They mainly moved alone or in pairs but did not have exclusive home ranges. Home range size tended to be larger during the dry season due to long movements to water sources. Birds generally preferred inselberg habitat and avoided farmland. During the dry season they additionally utilized gallery forests where water was readily available, and as a result of having to cross scrub savannah to get to water, scrub savannah was also more heavily used during the dry season. Birds bred between the late rainy (September–October) and the early dry season (November). Nest‐sites were associated with rocky boulders. Both sexes contributed to incubation. Daily egg survival rate calculated using the Mayfield method was 0.89 (0.83–0.95 95% confidence interval); no nests failed during the chick stage, but sample size was only four nests. Of all 14 nests found, 50% were depredated and only 29% of breeding attempts succeeded in producing Rock Firefinch chicks. A second breeding attempt was recorded when the first one failed. Chick production just about compensated adult mortality (measured in a separate study at the site) such that the population is probably stable at present. However, given the large uncertainty in our underlying assumptions, more data are needed to confirm this. We suggest that the presence of inselberg habitat in close proximity to water sources is the essential and limiting resource for this species.  相似文献   

19.
The diets of two sympatric, morphologically similar species of Moenkhausia were studied to investigate whether the manner in which they exploit food resources can help to explain their coexistence. Fish diets were evaluated during the low‐water season in six lakes of the upper Paraná River floodplain (Brazil). The results indicate that these two species segregate in relation to food resources and that the differential use of resources is probably a strategy that reduces competition during periods of a potential food shortage (the low‐water season).  相似文献   

20.
Riparian savanna habitats grazed by hippopotamus or livestock experience seasonal ecological stresses through the depletion of herbaceous vegetation, and are often points of contacts and conflicts between herbivores, humans and their livestock. We investigated how hippopotamus and livestock grazing influence vegetation structure and cover and facilitate other wild herbivores in the Mara region of Kenya. We used 5 km-long transects, each with 13 plots measuring 10 × 10 m2, and which radiate from rivers in the Masai Mara National Reserve and adjoining community pastoral ranches. For each plot, we measured the height and visually estimated the percent cover of grasses, forbs, shrubs and bare ground, herbivore abundance and species richness. Our results showed that grass height was shortest closest to rivers in both landscapes, increased with increasing distance from rivers in the reserve, but was uniformly short in the pastoral ranches. Shifting mosaics of short grass lawns interspersed with patches of medium to tall grasses occurred within 2.5 km of the rivers in the reserve in areas grazed habitually by hippos. Hence, hippo grazing enhanced the structural heterogeneity of vegetation but livestock grazing had a homogenizing effect in the pastoral ranches. The distribution of biomass and the species richness of other ungulates with distance from rivers followed a quadratic pattern in the reserve, suggesting that hippopotamus grazing attracted more herbivores to the vegetation patches at intermediate distances from rivers in the reserve. However, the distribution of biomass and the species richness of other ungulates followed a linear pattern in the pastoral ranches, implying that herbivores avoided areas grazed heavily by livestock in the pastoral ranches, especially near rivers.  相似文献   

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