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1.
Manipulations of herbivores in protected areas may have profound effects on ecosystems. We examine short‐term effects on tree species assemblages and resource utilization by a mesoherbivore and small‐size herbivores (ungulates <20 kg) in Sand Forest, after browsing release from a megaherbivore (elephant), or both a mega‐ and mesoherbivore (nyala), respectively. Effects were experimentally separated using replicated exclosures where all trees were counted, identified to species and browsing events recorded. Tree species assemblages were impacted by both elephant and nyala, and by each herbivore species individually. Tree turnover rates were higher where both herbivore species were present than in their combined absence. Diet was segregated among elephant, nyala and small‐size herbivores. Both resource specificity and browsing pressure by nyala increased in absence of elephant; small‐size herbivores increased resource specificity in absence of elephant, and increased browsing pressure in absence of both elephant and nyala. This implies interference competition with competitive release. The indirect effect of the manipulation of herbivore populations, through the removal of one or two herbivore species, caused a shift in tree species composition and diet of smaller‐size herbivores. These indirect effects, especially on tree species composition, can become critical as they affect vegetation dynamics, biodiversity and ecosystem processes. Therefore, in order to conserve habitats and biodiversity across all trophic levels, conservation managers should consider the effects of: (1) the full herbivore assemblage present; and (2) any effects of altering the relative and absolute abundance of different herbivore species on other herbivore species and vegetation.  相似文献   

2.
Elephant are considered major drivers of ecosystems, but their effects within small-scale landscape features and on other herbivores still remain unclear. Elephant impact on vegetation has been widely studied in areas where elephant have been present for many years. We therefore examined the combined effect of short-term elephant presence (< 4 years) and hillslope position on tree species assemblages, resource availability, browsing intensity and soil properties. Short-term elephant presence did not affect woody species assemblages, but did affect height distribution, with greater sapling densities in elephant access areas. Overall tree and stem densities were also not affected by elephant. By contrast, slope position affected woody species assemblages, but not height distributions and densities. Variation in species assemblages was statistically best explained by levels of total cations, Zinc, sand and clay. Although elephant and mesoherbivore browsing intensities were unaffected by slope position, we found lower mesoherbivore browsing intensity on crests with high elephant browsing intensity. Thus, elephant appear to indirectly facilitate the survival of saplings, via the displacement of mesoherbivores, providing a window of opportunity for saplings to grow into taller trees. In the short-term, effects of elephant can be minor and in the opposite direction of expectation. In addition, such behavioural displacement promotes recruitment of saplings into larger height classes. The interaction between slope position and elephant effect found here is in contrast with other studies, and illustrates the importance of examining ecosystem complexity as a function of variation in species presence and topography. The absence of a direct effect of elephant on vegetation, but the presence of an effect on mesoherbivore browsing, is relevant for conservation areas especially where both herbivore groups are actively managed.  相似文献   

3.
Sand Forest in the Maputaland region of KwaZulu-Natal in South Africa is deemed the most valuable, but also probably the most complex vegetation type of this part of the Maputaland–Pondoland–Albany hotspot of biodiversity. However, Sand Forest is under threat from the current human population growth in that region as well as from uncontrolled increases in wild herbivore numbers in conservation areas. The present study compares the impacts of herbivores and humans on the state of woody resources between two sites under differing utilisation regimes. Sand Forest was found to be a complex association of tree assemblages defined by different canopy and subcanopy properties. Although marked differences in the abundance of selected species were noted at the two sites, the Sand Forest remained dominated by fine-grained species under both utilisation regimes. The fine-grained nature of Sand Forest implies that regeneration depends on the creation of small canopy gaps either by natural processes, humans or elephants, while the creation of large gaps could transform it into woodland. Management of conservation areas where Sand Forest occurs should therefore concentrate efforts on regulating animal populations to levels that provide gap properties that favour forest regeneration.  相似文献   

4.
Sustained elephant browsing and intense burning could result in the loss of woodlands under conditions where elephant densities are high, such as in northern Botswana. Three woodland types dominated by Acacia erioloba, Baikiaea plurijuga and Colophospermum mopane were monitored in plots and contemporary recruitment rates of woody plants were compared with the associated local elephant densities and fire occurrences. Woodland types differed with respect to structure, extent of elephant damage and the occurrence of fire. Canonical correlations indicated that high extent of fire damage and high elephant densities did not covary within the woodland types investigated. Low tree densities in some woodland types were associated with high elephant densities and new elephant damage to plants increased with high elephant densities during the dry season. Plots with an apparent high fire frequency had lower tree densities and higher cover abundance of shrubs and seedlings.The annual rates of tree recruitment/loss in each woodland type were estimated through a model based on observed seedling recruitment, mortality and reversal to lower height classes due to combinations of fire occurrence and elephant browsing. The model suggested that elephants induce tree loss in woodlands dominated by plant species which are principal food sources. Fire however, seems to have a widespread effect across woodlands which could result in extensive tree loss.  相似文献   

5.
The substantial increase in elephant populations across many areas in southern Africa over past decades is prompting concerns about the effects on biodiversity. We investigated the outcomes of elephant disturbance on tree-species presence, density, and richness, and on alpha and beta diversity within riparian woodland in Chobe National Park, Botswana. We enumerated all tree species occurring in 32 plots (0.06 ha) along the Chobe riverfront. Plots were stratified by soil type (nutrient-rich alluvium vs. nutrient-poor Kalahari sand covering alluvium) and elephant impact (high vs. low impact on both soil types). We tested four predictions: elephants reduce tree density, richness, and alpha diversity; beta diversity is greater in vegetation subjected to high elephant impact; elephant impact on tree-species composition is greater on nutrient-poor than on nutrient-rich soil; and the loss or decline of abundant tree species on heavily disturbed sites is offset by an increase in abundance of functionally similar species, ones that are minor on lightly disturbed sites. Elephant browsing substantially affected tree-species composition, reducing density, species richness, evenness, and alpha diversity but had no effect on beta diversity. The dominant species on relatively undisturbed areas were partly replaced by functionally similar species on heavily disturbed sites. Soil type influenced species composition on lightly disturbed sites but was less important at higher elephant densities. Our findings are important for areas with extreme dry-season densities of elephants but should not be extrapolated to infer purported effects of elephants on tree diversity at lower densities.  相似文献   

6.
Seed predation and seedling mortality can act as strong demographic “bottlenecks” to sapling recruitment in African savanna woodlands. Fire also limits tree recruitment from saplings by suppressing their growth. I conducted field experiments with 13 woody plant species to assess the effects of seed burial on seedling emergence rates and effects of fire on seedling and sapling survival and growth rates over a period of 8 years at a savanna plot in central Zambia, southern Africa. Seed removal rates by small rodents varied among years and buried seeds had significantly higher emergence rates than seeds exposed to predators in most but not all the species. Annual burning reduced sapling growth in some species but in other species saplings experienced successive shoot die back even in the absence of fire. The findings show that for some woody species, seed predation is an important constraint to seedling recruitment but not for others and annual fires are important hindrances to demography and growth for some species but not others. Thus, demographic “bottlenecks” occur at different life history stages in different savanna woody species and these have the potential to alter woody tree competitive relationships and ultimately savanna structure.  相似文献   

7.
Elephants have a major influence on vegetation structure, composition and ecosystem processes, and are primary agents of habitat change in Africa. At moderate‐to‐high population densities, elephants can damage vegetation, especially when enclosed in protected areas. This study examines the effects of elephant browsing on woody trees in Majete Wildlife Reserve (WR), south‐western Malawi. Regression analysis is used to assess the associations of six factors known to drive elephant browsing in other areas and determine which ones have the most influence on browsing at Majete WR. Twenty‐four per cent of tagged trees had been subject to elephant browsing. The model with vegetation type, stem diameter and distance from permanent water correctly predicted browsing for 80% of the observations. Elephants mostly favoured riparian woodlands, followed by Acacia‐dominated woodland and Brachystegia‐dominated woodland. Browsing occurrence was negatively related to distance from permanent water and diameter at breast height(DBH). A larger number of trees, sampled at random and covering a larger portion of the reserve would provide more reliable estimates of browsing and related factors. Knowledge of time‐ and site‐specific factors affecting elephant browsing can be used to forecast future habitat transformations and manipulate the range of the elephants within the reserve.  相似文献   

8.
The reintroduction of African elephants into fenced game reserves throughout South Africa has presented managers with several challenges. Although elephants are a natural part of southern African ecosystems, their confinement to fenced protected areas in South Africa has exacerbated their potential to impact their habitats negatively. However, many studies investigating the impact of elephants have failed to control for the effects of other browsers on the vegetative community. In this study, we used location data on an elephant herd to delineate high-use and low-use areas. This paired design allowed us to minimize confounding factors that could explain differences in the structure, diversity and utilization of woody species. We found little evidence to suggest elephant-mediated change in, or selection for, the structure or diversity of woody species; however, our results suggest that elephants may be altering the composition of species by preferentially using areas with higher canopy diversity and by enhancing sapling recruitment. Although stripping of bark was higher in high-use areas, there was no evidence of differential mortality of tree species. Therefore, in our study area, and over the current time scale, elephants are having a negligible impact on the vegetative community.  相似文献   

9.
The ongoing loss of large trees and densification of shrubs are two prevalent processes that take place in African savannas, with profound consequences for their structure and function. We evaluated herbivore impacts on savanna woody communities using a long-term exclosure experiment in the Kruger National Park, South Africa, with three treatments: the exclusion of large mammals only (i.e. elephant and giraffe), exclusion of all herbivores larger than a hare, and areas open to all herbivores. We asked three questions: (1) How did variable exclusion of herbivores affect woody density and structure across the catena (i.e. riparian, sodic and crest vegetation)? (2) Did the exclusion of herbivores result in unique woody species composition? (3) Did herbivore exclusion result in a higher proportion of palatable species? After 17 years, we found that herbivores mainly affected the heights and densities of existing species, rather than leading to turnover of woody species assemblages. Although densities of individuals increased in the full exclosure (350 ha−1), the change was more moderate than expected. By contrast, mixed mega-and meso-herbivores decreased the number of trees and shrubs (decreases of 780 ha−1) via a variety of physical impacts. Meso-herbivores alone, on the other hand, had less impact on individual density (i.e. no change), but limited average height growth and canopy dimensions in certain habitat types. Where elephants are present, they are effective at reducing the density of woody stems to the point of counteracting woody encroachment, but at the same time are actively preventing the persistence of large trees (>5 m) as well as preventing trees from recruiting to larger size classes. However, the lack of massive recruitment and woody cover increases with elephant exclusion, especially for more preferred species, suggests that factors beyond elephants, such as dispersal limitation, seed predation, and drought, are also acting upon species.  相似文献   

10.
African forest elephants (Loxodonta cyclotis) are ecosystem engineers that browse and damage large quantities of vegetation during their foraging and movement. Though elephant trail networks and clearings are conspicuous features of many African forests, the consequences of elephant foraging for forest structure and diversity are poorly documented. In this study in northeastern Gabon, we compare stem size, stem density, proportional damage, species diversity, and species relative abundance of seedlings and saplings in the vicinity of seven tree species that produce elephant-preferred fruits (“elephant trees”) relative to control trees that do not. Across 34 survey trees, with a combined census area of 2.04 ha, we recorded data on 26,128 woody stems in three sizes classes. Compared with control trees, the area around elephant trees had the following: (a) a significantly greater proportion of damaged seedlings and a marginally greater proportion of damaged saplings (with 82% and 24% greater odds of damage, respectively); (b) no significant difference in stem density or species diversity; and (c) a significantly greater relative abundance of seedlings of elephant tree species. Increasing distance away from focal elephant trees was associated with significantly reduced sapling stem damage, significantly increased sapling stem density, and significantly increased sapling species diversity. Considered in sum, our results suggest that elephants can affect the structure and diversity of Afrotropical forests through their foraging activities, with some variation based on location and plant size class. Developing a more complete understanding of elephants’ ecological effects will require continued research, ideally with manipulative experiments.  相似文献   

11.
Effects of large mammalian herbivores on woody vegetation tend to be heterogeneous in space and time, but the factors that drive such heterogeneity are poorly understood. We examined the influence of fine‐scale habitat heterogeneity on the distribution and browsing effects of two of the largest African terrestrial mammals, the elephant and giraffe. We conducted this study within a 120‐ha (500 x 2,400 m) ForestGEO long‐term vegetation monitoring plot located at Mpala Research Center, Kenya. The plot traverses three distinct topographic habitats (“plateau,” “steep slopes,” and “valley”) with contrasting elevation, slope, soil properties, and vegetation composition. To quantify browsing damage, we focused on Acacia mellifera, a palatable tree species that occurs across the three habitat categories. Overall tree density, species richness, and diversity were highest on the steep slopes and lowest on the plateau. Acacia mellifera trees were tallest and had the lowest number of stems per tree on the steep slopes. Both elephant and giraffe avoided the steep slopes, and their activity was higher during the wet season than during the dry season. Browsing damage on Acacia mellifera was lowest on the steep slopes. Elephant browsing damage was highest in the valley, whereas giraffe browsing damage was highest on the plateau. Our findings suggest that fine‐scale habitat heterogeneity is an important factor in predicting the distribution of large herbivores and their effects on vegetation and may interact with other drivers such as edaphic variations to influence local variation in vegetation structure and composition.  相似文献   

12.
Abstract: Large herbivores such as elephants (Loxodonta africana) apparently have a negative impact on woody vegetation at moderate to high population densities. The confounding effects that fire, drought, and management history have may complicate assignment of such impacts to herbivory. We reviewed 238 studies published over 45 years and conducted a meta-analysis based on 21 studies that provided sufficient information on response of woody vegetation to elephants. We considered size and duration of studies, elephant densities, rainfall, fences, and study outcomes in our analysis. We detected a disproportionate citation of 20 published studies in our database, 15 of which concluded that woody vegetation responded negatively to elephants. Our analysis showed that high elephant densities had a negative effect on woody vegetation but that rainfall and presence of fences influenced these effects. In arid savannas, woody vegetation always responded negatively to elephants. In transitional savannas, an increase in elephant densities did not influence woody vegetation response. In mesic savannas, negative responses of woody vegetation increased when elephants occurred at higher densities, whereas elephants confined by fences also had more negative effects on woody plants than elephants that were not confined. Our analysis suggested that rainfall and fences influenced elephant density related impact and that research results were often site-specific. Local environmental conditions and site-specific objectives should be considered when developing management actions to curb elephant impacts on woody vegetation.  相似文献   

13.
The apparent influence of elephants on the structure of savannahs in Africa may be enhanced by management activities, fire and other herbivores. We separated the effect elephants have on grasses, woody seedlings (<0.5 m) and saplings (0.5–2 m) from the effect of tree canopies (canopy effect), and herbivory (park effect). We defined the canopy effect as the differences between plant abundances and diversity indices under tree canopies and 20 m away from these. Our testing of the park effect relied on the differences in the sub-canopy plant indices inside and outside a protected area that supported a range of herbivores. We based our assessment of the elephant effect on sub-canopy vegetation indices associated with elephant induced reductions in tree canopies. The park and canopy effects were more pronounced than the elephant effect. The park effect suppressed the development of woody seedlings into saplings. Conditions associated with tree canopies benefited woody plants, but not the grasses, as their indices were lower under trees. Elephants reducing canopies facilitated grass species tolerant of direct solar radiation. We concluded that management should consider other agents operating in the system when deciding on reducing the impact that elephants may have on vegetation.  相似文献   

14.
Elephants (Loxodonta africana) significantly alter ecosystem structure and composition through browsing (e.g. pollarding, debarking and toppling). Such browsing is predicted to intensify during severe drought which may become more common with climate change. Here, we make use of an elephant impact survey from 2012 to 2015 and during the El Nino drought of 2015–2016 at Pongola Game Reserve (107 km2), KwaZulu-Natal, to investigate how severe drought influenced damage severity of different tree heights and species by elephants in this small reserve. Contrary to expectations, damage to common species did not change with severe drought. Crown damage had the highest predicted probability across heights (29%–90%) and species (46%–75%) regardless of drought. However, we found severe drought increased the predicted probabilities of crown damage to smaller trees <4 m, mortality >6 m and severe damage at 4–6 m. Consequently, elephant damage during severe drought may alter vegetation structure by severely damaging or killing large trees (>4 m) and extensively damaging the crowns of trees <4 m. Long-term monitoring of elephant effects on woody vegetation is essential to enable science-based management in response to future drought and elephant damage (e.g. range expansion, beehive deterrents) to protect elephants and conserve woody vegetation.  相似文献   

15.
African elephant and black rhinoceros – both megaherbivores – impact negatively on the abundance of succulent plants, including tree succulents, in South Africa's subtropical thicket. We sampled 35 sites in subtropical thicket to assess historical and contemporary impacts of African elephant and black rhinoceros on the population structure of three species of succulent tree euphorbia. Population age structures were highly variable and showed no species-specific effects. Almost half the sites had growing populations dominated by young individuals. Sites having contemporary impacts had significantly fewer individuals in the 30–75 years age range, but this effect declined with increasing terrain slope. Eighty-one percent of sites assumed to have been impacted by megaherbivores historically had individuals that predated rhino and elephant extirpation in the region. This and other population age structure data indicate that megaherbivores and tree euphorbias coexisted historically on terrain of low relief. However, in areas not subject to contemporary impacts, euphorbia populations are likely to be much higher now than historically, owing to population relaxation after the extirpation of megaherbivores in the mid 1800s. Given the sensitivity of tree euphorbias to megaherbivore impacts, managers should consider monitoring the population structure of these species in order to identify thresholds of potential concern regarding megaherbivore impacts.  相似文献   

16.
Walter Leuthold 《Oecologia》1978,35(2):241-252
Summary Data on food habits and habitat preferences of four browsing herbivores (black rhinoceros, giraffe, gerenuk and lesser kudu) were analyzed to assess niche width for each species and niche overlap between pairs of species. All four species depended heavily on woody plants as food, and overlap in the utilization of different plant types (trees and shrubs, herbs, grasses, etc.) was very great in three of six species paris. When individual plant species were considered, markedly less overlap was apparent. Three of the four ungulate species preferred the most densely wooded vegetation type. Overlap in habitat preferences tended to be least in those pairs of species with the greates dietary overlap, which resulted in some degree of ecological separation. This was further increased by differences in browsing level. Seasonal variations in the browsing level of the giraffe had the effect of reducing overlap with the other species in the dry season, when food was in relatively short supply. Whether or not actual competition existed among the four ungulate species could not be established; in any event, it would probably be less important than possible competition exerted by the elephant, the dominant herbivore by far in the ecosystem. The ecological separation evident among the four browsing species probably permitted them to coexist in the area before the elephant reached its present dominant position and started altering the original vegetation.Formerly Tsavo East Research Station, Voi, Kenya  相似文献   

17.
Control of competing vegetation is recommended to ensure successful Yellow birch (Betula alleghaniensis Britton) regeneration within juvenile stands that do not sustain high enough sapling densities of this species. Four contrasting vegetation control treatments were tested to determine their effect on the growth and vigor of eight‐year‐old B. alleghaniensis saplings regenerating after final cutting of a shelterwood seed cut. Vegetation control treatments were TC (total circular removal), PC (circular removal of codominant competing vegetation), TS (total semicircular removal on 180° section), and NC (no vegetation control). Two years after treatment application, diameter growth significantly improved in response to vegetation control treatments, whereas sapling height growth did not. This pattern of biomass allocation was directly related to sapling etiolation, which increased with decreasing severity of vegetation removal. As a result, application of vegetation control, especially TC and PC treatments, was valuable in reducing signs of stress in saplings. However, increasing the severity of vegetation removal also made saplings more conspicuous to herbivores, which increased browsing, especially in the TC and PC treatments. Browsing was sufficient in some plots of the TC and PC treatments to overcome the vigor and diameter growth enhancements observed when browsing was negligible. In contrast to the TC and PC treatments, the TS treatment kept browsing very low while largely removing competition. The results suggest that B. alleghaniensis saplings established after final cutting of a shelterwood seed cut do take advantage of vegetation control treatments, but the decision to apply these treatments must include consideration of local herbivore population densities.  相似文献   

18.
This study evaluated the effects of humans and herbivores on woody vegetation structure in the woodlands and forests of Maputaland. The woody vegetation structure of plant communities at three sites within a similar environment but under different utilisation regimes since the early 1990s was evaluated by means of a stem diameter and tree height class analysis. The effects of human utilisation were evaluated on the land of the rural Manqakulane community; herbivore utilisation was evaluated in the Tembe Elephant Park; and because of the low levels of utilisation since 1992 the vegetation in the Tshanini Community Conservation Area was used as a benchmark for the comparisons. In both woodlands and sand forests, utilisation regime resulted in changes in stem diameter size class distributions, although the height structure remained unchanged. In communal land human-associated disturbance promoted the presence of small woody plants. In general, conserved land under animal utilisation, had the least small diameter woody plants, but most of the large trees, whereas in the absence of utilisation intermediate densities of small diameter woody plants were found but low densities of large trees. This study presents the first quantification of significant changes in the woody vegetation structure in Maputaland due to human utilisation, herbivore utilisation, or lack of utilisation. It also provides a timeframe within which land use can cause significant changes in vegetation structure.  相似文献   

19.
Porensky LM  Veblen KE 《Oecologia》2012,168(3):749-759
Spatial heterogeneity in woody cover affects biodiversity and ecosystem function, and may be particularly influential in savanna ecosystems. Browsing and interactions with herbaceous plants can create and maintain heterogeneity in woody cover, but the relative importance of these drivers remains unclear, especially when considered across multiple edaphic contexts. In African savannas, abandoned temporary livestock corrals (bomas) develop into long-term, nutrient-rich ecosystem hotspots with unique vegetation. In central Kenya, abandoned corral sites persist for decades as treeless ‘glades’ in a wooded matrix. Though glades are treeless, areas between adjacent glades have higher tree densities than the background savanna or areas near isolated glades. The mechanisms maintaining these distinctive woody cover patterns remain unclear. We asked whether browsing or interactions with herbaceous plants help to maintain landscape heterogeneity by differentially impacting young trees in different locations. We planted the mono-dominant tree species (Acacia drepanolobium) in four locations: inside glades, far from glades, at edges of isolated glades and at edges between adjacent glades. Within each location, we assessed the separate and combined effects of herbivore exclusion (caging) and herbaceous plant removal (clearing) on tree survival and growth. Both caging and clearing improved tree survival and growth inside glades. When herbaceous plants were removed, trees inside glades grew more than trees in other locations, suggesting that glade soils were favorable for tree growth. Different types of glade edges (isolated vs. non-isolated) did not have significantly different impacts on tree performance. This represents one of the first field-based experiments testing the separate and interactive effects of browsing, grass competition and edaphic context on savanna tree performance. Our findings suggest that, by excluding trees from otherwise favorable sites, both herbaceous plants and herbivores help to maintain functionally important landscape heterogeneity in African savannas.  相似文献   

20.
Rewilding and translocations of large herbivores for conservation purposes have increased in recent times, with numerous introductions inside and outside their native range. This study aims to analyze the use of threatened plant taxa as a possible ecological indicator of large herbivore introductions. We examined the effects of a threatened large ungulate, the Barbary sheep (Ammotragus lervia), on both endangered and vulnerable woody taxa after its introduction in 1970. Contrary to our hypothesis, the herbivore impact on threatened woody species was higher than that found on widespread woody plants. The results reveal that 35.7% of the threatened species showed the highest possible level of herbivore damage in contrast to 6.5% for the widespread species. Threatened species were preferred over common plants, probably due to their greater palatability. Overall plant cover, including neighboring species, was also an important factor determining browsing damage and, thus, habitats with low ground cover should be particularly considered in conservation plans. Herbivore damage on common taxa should be taken with caution since they could mask unsustainable herbivore densities for threatened woody taxa or protected habitats. The use of threatened woody taxa through the studied ecological indicators (herbivore damage, plant preferences, habitat use and regeneration success) represented a useful tool to assess the sustainability of large herbivores introductions and to establish a priority conservation ranking for threatened plant species. These findings highlight the deleterious effects of overabundant ungulate populations regardless its origin (exotic or native) and the need of monitoring threatened woody taxa to better estimate the suitability and sustainability of large herbivore introductions.  相似文献   

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