Contemporary and historical impacts of megaherbivores on the population structure of tree euphorbias in South African subtropical thicket

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.     

The flora of the Addo Elephant National Park, South Africa: are threatened species vulnerable to elephant damage?

The Addo Elephant National Park in the warm-temperate Eastern Cape was proclaimed in 1931 to protect one of the four elephant populations in South Africa which survived into the present century. However, since the late 1970s, a major objective of the Park is to protect intact a viable example of succulent thicket – the regional ecosystem. Succulent thicket is endemic to the Eastern Cape and forms the major component of the Albany Centre, a floristic region of high diversity and endemism, especially among succulent shrubs and geophytes. This ecosystem is poorly represented in the reserve system and is highly threatened by overgrazing and clearing for agriculture. Under the present stocking rate of more than two elephants/km², succulent thicket in the Park has been severely impacted: in particular, biomass, stature and plant diversity have been reduced. We show that the flora of the Park is of great regional significance. Of a total of 581 species, 12.4% were classified as regional endemics and/or Red Data Book species (collectively termed 'species of special concern), and 32.2% were not known to be conserved in any other succulent thicket protected area. Relative to the remainder of the flora, both categories of species were over-represented among succulent shrubs (predominantly Euphorbiaceae and Mesembryanthemaceae) and geophytes (predominantly Asphodelaceae and Hyacinthaceae). This taxonomic and biological profile coincides with that of species vulnerable to local extinction as a result of elephant impacts. The Park authorities will need to find a solution to the conflict between maintaining a large population of megaherbivores, and maintaining the structure and diversity of succulent thicket.     

The affects of artificial water availability on large herbivore ranging patterns in savanna habitats: a new approach based on modelling elephant path distributions

Aim  Artificial water points are often used in protected savanna ecosystems to maintain populations of large herbivores. However, these interventions lead to increased ranging and foraging pressure and can negatively impact important habitats and species. This study investigated the influence of artificial water provision on the foraging and movement paths of an African elephant population and modelled the impact of changing water availability on sensitive habitat types.
Location  Tembe Elephant Park (TEP), KwaZulu-Natal, South Africa.
Methods  We mapped and classified 414 km of elephant movement and foraging paths in a 300-km² fenced protected area. The data were analysed to determine the relationship between path size, distribution and distance to water. We also used a logistic modelling approach to explore the predicted effects of removing artificial water points on path distribution.
Results  Elephant paths were unevenly distributed throughout the habitats of TEP and the most established and heavily used paths were found closest to water. We also discovered a number of elephant 'rest areas' along the paths, which were distinct clearings that tended to be close to water and in sand forest habitat. Our model predicted that the removal of artificial water points would reduce the area crossed by elephant paths by 79%, leading to an 89% reduction in the presence of elephant paths in sand forest.
Main conclusions  Our study provides further evidence that manipulating surface water availability can be a useful tool for managing large herbivore impacts on vegetation and acts as the basis for further research on the trade-offs between conservation objectives.     

Relevance of elephant herbivory as a threat to Important Plants in the Addo Elephant National Park, South Africa

Although elephants are recognized as keystone species, the mechanisms of their impacts on biodiversity and community structure are rarely identified. In the Addo Elephant National Park (AENP), South Africa, elephant Loxodonta africana herbivory is apparently responsible for a significant reduction in plant richness, especially among the regionally rare and endemic small succulent shrubs and geophytes (Important Plants). We used faecal analysis to investigate the utilization of Important Plants in elephant diet in the AENP. Ninety plant species were identified in the diet. Only 14 of the 77 ( c . 18%) Important Plants previously thought particularly vulnerable to elephant browsing occurred in the diet, while at least 6% of species for which there are data were avoided. This refutes the generally held belief that elephant herbivory is the major driver of decline among Important Plants, and emphasizes the likely contribution of other mechanisms (e.g. knock-on effects, trampling, zoochory, etc.) to this phenomenon. The accurate prediction of impacts caused by elephants in the AENP and elsewhere, therefore requires an understanding of these previously marginalized mechanisms. By demonstrating appropriate cause-and-effect relationships between elephants and ecosystem change, we will be able to move beyond assuming that all the observed changes are due to elephant herbivory.     

Elephant both Increase and Decrease Availability of Browse Resources for Black Rhinoceros

Despite the apparent dual role of elephant Loxodonta africana in shaping the food niches of large herbivores, empirical studies focus on their role in facilitating foraging opportunities, while declining resource opportunities (a necessary requirement for competition) are rarely quantified. Our study investigates the relative importance of elephant in these processes by quantifying potential browsing opportunities (using total and preferred biomass, between‐bite distances and bite mass) for black rhinoceros Diceros bicornis along a gradient of elephant utilization in the succulent thickets of the Addo Elephant National Park, South Africa. We show that browse biomass and potential between‐bite harvest rates initially increase with the intensity of elephant utilization through the formation and spread of elephant pathways in otherwise impenetrable thicket. At the maximum, modeled estimates of total and preferred biomass are on average 223 percent and 254 percent higher, respectively, than that recorded in the absence of elephant (Exclosures); potential between‐bite harvest rates are 75 percent higher. With continued elephant utilization, however, browse biomass declines and between‐bite distances increase as the pathways expand and coalesce and canopy height declines. Our model of the change in potential browsing opportunities for rhinoceros reflects the accumulated effects of elephant over time, i.e., the effects accumulate until the relationship switches from increased to reduced availability. With this we demonstrate the key role of elephant for rhinoceros foraging, linked to a potential loss of this role at higher levels of utilization.     

Landscape dysfunction and reduced spatial heterogeneity in soil resources and fertility in semi‐arid succulent thicket,South Africa

Abstract Succulent thicket in the Eastern Cape of South Africa is one of many rangeland ecosystems in the world that displays evidence of unsustainable grazing pressure. Widespread transformation of succulent thicket has resulted in the replacement of the typical two‐phase perennial vegetation patches with a structurally simple field layer of ephemeral and weakly perennial grasses and forbs. We hypothesized that (i) transformation of succulent thicket leads to a switch from a spatially heterogeneous landscape to a homogenous and relatively infertile state; and (ii) that this loss of fertility is associated with a breakdown in the processes that conserve resources and promote water use efficiency. We tested these hypotheses at five fenceline contrast sites in Sundays River Thicket, an arid form of succulent thicket. We compared soil fertility (organic carbon, available nitrogen and phosphorus), texture, matric potential, and surface microtopography in the two main microhabitats on either side of the fencelines using a nested anova. Our results show that intact Sundays River thicket has a distinct spatial pattern of soil fertility where nutrients and organic carbon are concentrated under the patches of perennial shrubs, compared with under canopy trees. Transformation results in a significant homogenization of this pattern and an overall reduction in the fertility of the landscape. The proportion of the landscape surface that promotes infiltration due to a distinct litter layer decreases from 60% to 0.6%. Soil moisture retention (matric potential) also decreases with transformation. We interpret these patterns within the framework of semi‐arid landscape functionality.     

金沙江干旱河谷植物群落的数量分类及其结构分异的环境解释

植物群落的空间分异格局是异质生境条件下物种性状、种间相互作用等生态学过程共同作用的结果, 对其分析有助于深入理解群落构建进程。本文基于金沙江流域干旱河谷116个样点562个样方的植物群落调查数据, 采用自适应仿射传播聚类的方法进行群落数量分类, 运用莫兰特征向量地图, 和方差分解的方法对影响群落结构的空间和环境因子进行分析。结果表明: (1)自适应仿射传播聚类将金沙江干旱河谷的植物群落分为30组, 可归为7个植被型, 23个群系, 以稀树草原(30.0%)、暖性落叶阔叶灌丛(55.7%)为最主要的植被类型。(2)年均温和干燥指数是限制金沙江干旱河谷植物群落分布的主要环境因子。稀树草原、肉质灌丛、常绿阔叶灌丛是典型的干热河谷植被类型; 暖性落叶阔叶灌丛、常绿硬叶林是干暖河谷植被的优势类型; 暖性针叶林、落叶阔叶林则主要在干温河谷环境占优势。(3)纯环境因子可以解释群落物种组成变化的5.5%, 纯空间因子可以解释的物种组成变化为22.5%, 有空间结构的环境因子部分为6.6%, 未解释的部分为65.4%。在诸多环境因子中, 年均温及干燥指数的不同显示了不同群落生境的重要差异, 并显著影响到群落的分布格局。大尺度的空间因子则主要通过地理隔离对群落结构的差异产生影响。     

水源圈研究进展

在干旱、半干旱区草场上,由于饮水的需求导致牲畜经常性地集中在水源点周围,从而使草地生态系统以水源点为中心呈梯度退化,形成水源圈.按照放牧影响程度的大小,以水源点为中心由内向外可将水源圈划分为“牺牲带”、过渡带和自然带3部分,水源圈内的植被、土壤等对放牧的响应也呈梯度变化.由于水源圈内存在着“天然的放牧梯度”,使其成为研究放牧对植被、土壤的影响和自然资源保护及其可持续利用的绝佳场所.本文对水源圈的概念、研究的发展历程和研究方法等进行了综述,并对国内研究现状进行了评述,以促进干旱、半干旱区草场的科学管理和可持续利用.     

水源圈研究进展

在干旱、半干旱区草场上,由于饮水的需求导致牲畜经常性地集中在水源点周围,从而使草地生态系统以水源点为中心呈梯度退化,形成水源圈.按照放牧影响程度的大小,以水源点为中心由内向外可将水源圈划分为“牺牲带”、过渡带和自然带3部分,水源圈内的植被、土壤等对放牧的响应也呈梯度变化.由于水源圈内存在着“天然的放牧梯度”,使其成为研究放牧对植被、土壤的影响和自然资源保护及其可持续利用的绝佳场所.本文对水源圈的概念、研究的发展历程和研究方法等进行了综述,并对国内研究现状进行了评述,以促进干旱、半干旱区草场的科学管理和可持续利用.     

Evaluating performance of aerial survey data in elephant habitat modelling

Aerial survey data are widely used to model distribution of wildlife. However, their performance in habitat modelling remains largely untested. We used aerial survey and satellite‐linked Global Positioning System (GPS) collar data for elephants, to test (i) whether there is an optimal spatial resolution of predictor variables at which habitat models based on aerial survey data that are uncorrected for locational error can accurately predict elephant habitat and (ii) whether habitat models based on these data sets can accurately predict the presence of elephants in closed woodland habitats. We applied maximum entropy modelling (Maxent) to these data sets and used the Normalised Difference Vegetation Index (NDVI) as well as distance from water points as the habitat predictors to answer these questions. Our results demonstrate better ability of aerial survey data to predict elephant presence at the coarser spatial resolution of 1000 m of both predictor variables. Habitat models derived from aerial survey data underpredicted elephant presence in more closed woodland habitats than those derived from GPS collar data. This result implies that elephants located under dense tree canopies are likely missed during an aerial survey. Our study is one of the first to empirically test and report results on the poor performance of aerial survey data in habitat modelling especially in dense woodlands.     

Punching above their weight: low-biomass non-native plant species alter soil properties during primary succession

Non‐native invasive plants can greatly alter community and ecosystem properties, but efforts to predict which invasive species have the greatest impacts on these properties have been generally unsuccessful. An hypothesis that has considerable promise for predicting the effects of invasive non‐native plant species is the mass ratio hypothesis (i.e. that dominant species exert the strongest effects). We tested this hypothesis using data from a four year removal experiment in which the presence of two dominant shrub species (one native and the other not), and subordinate plant species, were manipulated in factorial combinations over four years in a primary successional floodplain system. We measured the effects of these manipulations on the plant community, soil nutrient status and soil biota in different trophic levels of the soil food web. Our experiment showed that after four years, low‐biomass non‐native plant species exerted disproportionate belowground effects relative to their contribution to total biomass in the plant community, most notably by increasing soil C, soil microbial biomass, altering soil microbial community structure and increasing the abundance of microbial‐feeding and predatory nematodes. Low‐biomass, non‐native plant species had distinct life history strategies and foliar traits (higher foliar N concentrations and higher leaf area per unit mass) compared with the two dominant shrub species (97% of total plant mass). Our results have several implications for understanding species’ effects in communities and on soil properties. First, high‐biomass species do not necessarily exert the largest impacts on community or soil properties. Second, low‐biomass, inconspicuous non‐native species can influence community composition and have important trophic consequences belowground through effects on soil nutrient status or resource availability to soil biota. Our finding that low‐biomass non‐native species influence belowground community structure and soil properties more profoundly than dominant species demonstrates that the mass ratio hypothesis does not accurately predict the relative effects of different coexisting species on community‐ and ecosystem‐level properties.     

Effects of goat pastoralism on ecosystem carbon storage in semiarid thicket,Eastern Cape,South Africa

Abstract Intensive pastoralism with goats transforms semiarid thicket in the Eastern Cape, South Africa from a dense vegetation of tall shrubs to an open landscape dominated by ephemeral grasses and forbs. Approx. 800 000 ha of thicket (which prior to the introduction of goats had a closed canopy and a Portulacaria afra Jacq. component) have been transformed in this manner. Ecosystem C storage in intact thicket and loss of C due to transformation were quantified. Carbon storage in intact thicket was surprisingly high for a semiarid region, with an average of 76 t C ha^?1 in living biomass and surface litter and 133 t C ha^?1 in soils to a depth of 30 cm. Exceptional C accumulation in thicket may be a result of P. afra dominance. This succulent shrub switches between C₃ and CAM photosynthesis, produces large quantities of leaf litter (approx. 450 g m^?2 year^?1) and shades the soil densely. Transformed thicket had approx. 35% less soil C to a depth of 10 cm and approx. 75% less biomass C than intact thicket. Restoration of transformed thicket landscapes could consequently recoup more than 80 t C ha^?1.     

Changes in arthropod diversity along a land use driven gradient of shrub cover in savanna rangelands: identification of suitable indicators

Shrub encroachment linked to heavy grazing has dramatically changed savanna landscapes, and is a major form of rangeland degradation. Our understanding of how shrub encroachment affects arthropod communities is poor, however. Here, we investigate the effects of shrub encroachment on abundance and diversity of ground-dwelling (wingless) arthropods at varying levels of shrub cover in the southern Kalahari. We also ascertain if invertebrate assemblage composition changes with habitat structure and identify which aspects of habitat structure (e.g., grass cover, herbaceous plant cover, shrub density) correlate most strongly with these changes. Ant, scorpion and dung beetle abundance increased with shrub cover, whereas grasshoppers and solifuges declined. Spider and beetle abundance exhibited hump-shaped relationships with shrub cover. RTU richness within orders either mirrored abundances, or exhibited no trend. Shrub density was the habitat component most correlated with similarities between invertebrate assemblages. Ground-dwelling arthropods showed clear shifts in species assemblage composition at a similarity level of 65% according to shrub density. Changes in indicator species showed that within the Tenebrionidae (darkling beetles), certain species respond positively to shrub thickening, replacing other species within the Family. Small-bodied, wingless Scarabaeidae (dung beetles) tended to increase with increased shrub density and three species emerged as significant indicators of more thickened habitats, although this might be a response to greater dung availability, rather than habitat structure itself. We conclude that because ground-dwelling invertebrates showed such clear responses in species assemblage composition, they present excellent candidates for use as indicator species in further studies into bush encroachment.     

皖南黟县次生灌草丛生物量的研究

本文对地处中亚热带的安徽黟县次生灌丛和灌草丛以及常绿阔叶幼林共9个样地的生物量进行了研究。结果表明:黟县常绿阔叶幼林、次生灌丛和灌草丛的地上部分为81.74、16.83和5.28t/ha。所调查的5个群落类型的地上部分生物量和生产力可回归成线性方程Y=3.88+0.14X,净生产力随生物量的增加而提高。     

Heterogeneity in African savanna elephant distributions and their impacts on trees in Kruger National Park,South Africa

1. Though elephants are a major cause of savanna tree mortality and threaten vulnerable tree species, managing their impact remains difficult, in part because relatively little is known about how elephant impacts are distributed throughout space.
2. This is exacerbated by uncertainty about what determines the distribution of elephants themselves, as well as whether the distribution of elephants is even informative for understanding the distribution of their impacts.
3. To better understand the factors that underlie elephant impacts, we modeled elephant distributions and their damage to trees with respect to soil properties, water availability, and vegetation in Kruger National Park, South Africa, using structural equation modeling.
4. We found that bull elephants and mixed herds differed markedly in their distributions, with bull elephants concentrating in sparsely treed basaltic sites close to artificial waterholes and mixed herds aggregating around permanent rivers, particularly in areas with little grass.
5. Surprisingly, we also found that the distribution of elephant impacts, while highly heterogeneous, was largely unrelated to the distribution of elephants themselves, with damage concentrated instead in densely treed areas and particularly on basaltic soils.
6. Results underscore the importance of surface water for elephants but suggest that elephant water dependence operates together with other landscape factors, particularly vegetation community composition and historical management interventions, to influence elephant distributions.

     

云南西双版纳尚勇保护区亚洲象对栖息地的选择

以西双版纳国家级自然保护区尚勇子保护区内的亚洲象种群为研究对象, 利用村寨调查、样线调查和3S(GIS、GPS、RS) 技术对该地区野生亚洲象的栖息地状况和亚洲象对栖息地选择利用进行了初步的研究。结果表明亚洲象喜欢海拔1 000 m以下的区域, 坡度小于10°的区域, 坡位为平坦的沟谷和山坡的下部, 坡向为南、北两个方向。偏好的植被类型有竹阔混交林、灌丛和高山草甸。研究还发现尚勇保护区及周边区域亚洲象栖息地的丧失和日益增加的非法盗猎活动已经严重威胁到该地区野生亚洲象种群的生存。     

Responses of mammalian carnivores to land use in arid savanna rangelands

Shrub encroachment due to heavy grazing has led to dramatic changes in arid savanna landscape structure worldwide. It is considered to be one of the most threatening forms of rangeland degradation altering plant diversity. However, possible impacts of shrub encroachment on species diversity at higher trophic levels remain poorly understood. Additionally, indirect effects, such as changes of trophic interactions, are often ignored when trying to understand changes in biodiversity patterns. In this study, conducted in the southern Kalahari, we explored how the diversity of small carnivores and their prey is affected by shrub encroachment. We analysed the relationships between abundance and diversity of small carnivores, the availability of their main prey groups (coleopterans, termites, grasshoppers, rodents) and the structural diversity of the landscape (shrub, grass and herb cover). Eight hundred track surveys were conducted to determine carnivore abundance on 20 rangeland habitats, which represented a gradient of grazing intensity. Prey availability was surveyed for each study site and related to vegetation cover.

Results show a significant impact of shrub cover on abundance and diversity of carnivores and their prey. The diversity of both, carnivores and their prey, showed a hump-shaped response to increasing shrub cover whereas relative carnivore abundance decreased. Availability of prey groups was affected differently by shrub cover increase. Diversity of carnivores was best predicted by shrub cover (R²>0.7, p<0.001) indicating the overriding role of habitat structure as compared to prey availability. We conclude that intermediate shrub cover values enrich structural diversity of savanna landscape and in consequence sustain diversity of small carnivores and their prey in arid and semiarid ecosystems.     

Post-dispersal seed predation of an exotic weed,Mimosa pigra L., in the Northern Territory

The removal of seeds of the exotic shrub Mimosa pigra from the ground beneath a thicket of the plant and from the open floodplain surrounding the thicket was studied experimentally. Seeds placed beneath the canopy of the M. pigra thicket were taken predominantly by vertebrates (probably omnivorous rodents) while those on the open floodplain were taken predominantly by ants. After 14 days exposure with no barriers to seed removal, less than half of the seeds placed in either habitat had disappeared. Of the seeds that were removed, over 75% had disappeared from both habitats within 5 days. It is unlikely that post-dispersal seed predation appreciably inhibits the spread or maintenance of stands of this weed in tropical Australia.     

Rooting characteristics of two widely distributed woody plant species growing in different karst habitats of southwest China

A series of studies claimed that deep root development of plant established in karst regions was facilitated by fractured bedrock beneath the shallow soils; however, bedrock is not a uniform medium for root proliferation. We hypothesized plant species that survived in different karst habitats had some other rooting characteristics rather than deep penetration. To test the hypothesis, coarse root systems of two widely distributed woody species (one tree and one shrub) growing in three typical rocky karst habitats (shallow soil, loose rocky soil and exposed rock) were excavated in karst region of southwest China. Root systems were investigated based on four parameters: maximum rooting depth, maximum radial extent, root tapering pattern and root curvature. In all the three habitats, maximum rooting depths were no deeper than 120 and 40 cm for the tree and shrub species, respectively. Maximum radial extents were extremely large compared with maximum rooting depth, indicating that rooting characteristics were dominated by horizontal extension rather than deep penetration. Roots of both species growing in shallow soil habitat tapered gradually and curved slightly, which was consistent with the specific characteristics of this habitat. On the contrary, roots of the tree species growing in the other two habitats tapered rapidly but curved slightly, while roots of the shrub species tapered gradually but curved strongly. It was speculated that limited depths and rapid tapering rates of the tree roots were likely compensated by their utmost radial extensions, while the shrub species might benefit from its root curvature as the associated root tropisms may increase the ability of root to encounter more water and contribute to potentially high resource absorption efficiency. Our results highlight the importance of taking shallow-rooted species into account in understanding the distribution of natural plant communities and predicting future vegetation dynamics in karst regions.     

Shift in Black Rhinoceros Diet in the Presence of Elephant: Evidence for Competition?

In African large herbivore assemblages, megaherbivores dominate the biomass and utilise the greatest share of available resources. Consequently, they are considered a separate trophic guild that structures the food niches of coexisting large herbivores. However, there exists little empirical evidence on how food resources are shared within this guild, and none for direct competition for food between megaherbivores. Using the histological analysis of faeces, we explore this phenomenon for African elephant Loxodonta africana and black rhinoceros Diceros bicornis in the Addo Elephant National Park, South Africa, where the accumulated impacts of elephant have reduced browse availability. Despite being unable to generalise beyond our study sites, our observations support the predictions of competition theory (as opposed to optimality theory) by showing (1) a clear seasonal separation in resource use between these megaherbivores that increased as resource availability declined, and (2) rhinoceros changed their selectivity in the absence of elephant (using an adjacent site) by expanding and shifting their diet along the grass-browse continuum, and in relation to availability. Although black rhinoceros are generally considered strict browsers, the most significant shift in diet occurred as rhinoceros increased their preferences for grasses in the presence of elephant. We speculate that the lack of specialised grazing adaptations may increase foraging costs in rhinoceros, through reduced harvest- and handling-efficiencies of grasses. In the short-term, this may be off-set by an enhanced tolerance for low quality food and by seasonally mobilising fat reserves; however, the long-term fitness consequences require further study. Our data suggest that managing elephant at high densities may compromise the foraging opportunities of coexisting browsers. This may be particularly important in small, fenced areas and overlapping preferred habitats where impacts intensify.