Predicting the habitat usage of African black rhinoceros (Diceros bicornis) using random forest models

Species distribution models are often used in ecology to ascertain relationships between environmental variables and species presence. Modelling to understand this relationship can be used to aid conservation management strategies. In this paper, we applied the random forest classification method to predict habitat used by black rhino for browsing. The random forest model was created using detailed habitat data collected from Ol Pejeta Conservancy in Kenya. Variables from plots where rhino had been present were compared to those not used by rhino. Independent data were used to test the predictive accuracy of the rules generated. The model performed well with the independent test data, correctly classifying 69% of the sampling plots where black rhino were present. Important habitat features that affected rhino presence were browse availability and density of vegetation, with Vachellia drepanolobium (formerly Acacia) and Euclea divinorum being important components. The analysis also highlighted areas of potential high browse pressure, which should be the focus of continued monitoring and management.     

Fine‐scale habitat heterogeneity influences browsing damage by elephant and giraffe

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.     

Effects of browse availability and quality on black rhino (Diceros bicornis michaeliGroves 1967) diet in Nairobi National Park,Kenya

Investigations were conducted to study the effects of browse availability and quality on black rhinoceros (Diceros bicornis michaeliGroves 1967) diet in Nairobi National Park, Kenya, between September 1994 and February 1995. Line intercept method was used to assess availability of browse. The ratios of browsed crowns to the total crown were applied in the estimate of browse use. Proximate analysis and extraction methods were used, respectively, to appraise browse nutritional quality and phytochemical status of twelve species. Thirty‐four plant species were identified as potential black rhino browse, whose availability differed significantly (P < 0.001). Lippia javanica (20.581 ± 3.101, n = 544) and Lannea cornuta (0.963 ± 0.529, n = 544) were the most and least available species, respectively. Only 32 species, however, were utilized, as L. cornuta and Rhynchosia hirta did not show any level of rhino feeding activity. Browse use was found to vary significantly (P < 0.001) during the study period. Grewia similis was most (20.564 ± 2.764, n = 544) and Commelina africana least (0.974 ± 0.974, n = 544) exploited, respectively. Differences in total alkaloids and phenols, crude fibre, crude protein and total ash contents among species investigated were highly significant (P < 0.001). Ether extract values were similar (P= 0.0933) across plant taxa. The study shows that the black rhino is a selective browser feeding on plants with low phenol and alkaloid contents and high fibre values. Of the nutritionally similar species, browsing was driven by availability. These findings suggest that a key habitat factor important in black rhinoceros conservation include diverse plant species with low levels of phenols and alkaloids.     

The savanna ecology of Kidepo Valley National Park

(1) Food habits of elephant and giraffe were recorded in short sample periods over several years using the feeding minutes technique. (2) Twelve species of common plants were collected on these occasions, separated into their main parts and analysed chemically. (3) Damage to trees was also recorded at intervals. (4) Fifty-nine species of plant were eaten by elephant. During wet months their diet was predominantly grass but during dry months woody vegetation became more important. Giraffe were observed eating from thirty-nine species of plant which were almost entirely browse throughout the year. At least eighteen of these species were also eaten by elephant. (5) Three of the browse species most important to giraffe were also important to elephant in dry seasons. In the driest month the species eaten by giraffe formed more than two-thirds of the observed diet of elephants. (6) Chemical analyses of plants show that browse leaves had a higher level of protein, fat and minerals than most other plants. (7) The trunk of an elephant and the tongue of a giraffe are highly sensitive organs which permit food selection. Elephant in particular appeared to alter their food habits seasonally and it is suggested that this may be an attempt to obtain a balanced intake of nutrients. (8) Due to the effect of elephant and other agencies there is an accelerating decline in Acacia gerrardii woodlands. Browsing and fire are independently preventing their regeneration. (9) The extensive overlap of food requirements on preferred but diminishing browse resources implies potential if not already existing competition between elephant and giraffe. (See Part III.)     

Not all ants are equal: obligate acacia ants provide different levels of protection against mega‐herbivores

In obligate ant–plant mutualisms, the asymmetric engagement of a single plant species with multiple ant species provides the opportunity for partners to vary in their behaviour. Variation in behaviour has implications for the interactions with third‐party species such as herbivores. This study assessed the effect of obligate ant‐mutualists (Crematogaster mimosae, Crematogaster nigriceps and Tetraponera penzigi) inhabiting the African ant‐acacia (Acacia drepanolobium) on three mega‐herbivore browsers: the Maasai giraffe (Giraffa camelopardalis tippelskirchi), the reticulated giraffe (Giraffa c. reticulata) and the black rhino (Diceros bicornis). Giraffes are abundant and wide‐ranging herbivores of the acacias, whereas black rhinos are localized and perennial herbivores of the acacias. Multiyear field studies comparing the ants’ aggressive behaviour and browsing by mega‐herbivores suggested differences between the tending abilities of the primary ant species inhabiting A. drepanolobium. Trees occupied by the aggressive ant species C. mimosae had significantly less browsing by giraffes and black rhino than trees occupied by other ant species. The results of this study provide evidence that ant‐mutualists on African acacias can serve as deterrents to mega‐herbivores and that different ant species vary in their tending abilities.     

Introduction of giraffe changes acacia distribution in a South African savanna

Large mammal herbivores can have significant effects on the structure and composition of plant communities. We studied the impacts of an introduced giraffe population on Acacia species at Ithala Game Reserve in South Africa. Browse intensity and Acacia mortality were assessed in field transects, and in road transects covering the reserve road network. Several Acacia species occurring in high‐density giraffe areas had high levels of mortality. Populations of Acacia davyi were extinct in areas accessible to giraffe. Most A. caffra trees within giraffe browse height were dead and A. karroo, the most common species, was also heavily affected. Some species, including A. tortilis, showed no or very low mortality attributable to giraffe browsing. Healthy populations of sensitive species occurred in areas within, and adjacent to, the reserve in areas with low or no giraffe browsing. Areas too steep for giraffe access formed spatial refuges for these trees. The differential mortality that is occurring as a consequence of giraffe browsing is altering species composition and species distribution in this savanna landscape.     

Food habits of giraffe in Tsavo National Park, Kenya

In the context of a broader ecological investigation, food habits of giraffe were studied in Tsavo National Park (East). The only method employed was direct observation of feeding animals in the field. Each instance in which one animal fed on one individual plant was counted as one food record for this plant species; 4025 records are analysed. A total of sixty-six plant species was found to be eaten, the great majority being trees and shrubs, with a few creepers and vines. There were marked seasonal differences in the diet of giraffe, deciduous trees, shrubs and vines being dominant in the green season, evergreen plants (partly in riverine forest) in the dry season. All the trees and larger shrubs common in the study area were eaten by giraffe, while few records for very small shrubs and none for herbs and grasses were obtained. An analysis of the available vegetation was made in part of the dry-season habitat, and for twenty species the frequency in the habitat was compared with the frequency in the giraffe's diet, revealing selection for or against certain species. Giraffe utilized the upper vegetation layers, where available, but overall c. 50% of all browsing was below 2 m above ground, i.e. within reach of smaller browsers. Results of this study are compared with what is known on food habits of giraffe in other areas. Possible competition of giraffe with other browsers and the relationship between giraffe and their habitat are discussed. Continued survival of giraffe, and other browsers, in Tsavo National Park depends primarily on (1) adequate control of fire, and (2) the impact of future vegetation changes on the amount and variety of available browse plants.     

Ecological separation among browsing ungulates in Tsavo East National Park,Kenya

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     

Elephant herbivory in Majete Wildlife Reserve,Malawi

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.     

Standardising Home Range Studies for Improved Management of the Critically Endangered Black Rhinoceros

Comparisons of recent estimations of home range sizes for the critically endangered black rhinoceros in Hluhluwe-iMfolozi Park (HiP), South Africa, with historical estimates led reports of a substantial (54%) increase, attributed to over-stocking and habitat deterioration that has far-reaching implications for rhino conservation. Other reports, however, suggest the increase is more likely an artefact caused by applying various home range estimators to non-standardised datasets. We collected 1939 locations of 25 black rhino over six years (2004–2009) to estimate annual home ranges and evaluate the hypothesis that they have increased in size. A minimum of 30 and 25 locations were required for accurate 95% MCP estimation of home range of adult rhinos, during the dry and wet seasons respectively. Forty and 55 locations were required for adult female and male annual MCP home ranges, respectively, and 30 locations were necessary for estimating 90% bivariate kernel home ranges accurately. Average annual 95% bivariate kernel home ranges were 20.4 ± 1.2 km², 53 ±1.9% larger than 95% MCP ranges (9.8 km² ± 0.9). When home range techniques used during the late-1960s in HiP were applied to our dataset, estimates were similar, indicating that ranges have not changed substantially in 50 years. Inaccurate, non-standardised, home range estimates and their comparison have the potential to mislead black rhino population management. We recommend that more care be taken to collect adequate numbers of rhino locations within standardized time periods (i.e., season or year) and that the comparison of home ranges estimated using dissimilar procedures be avoided. Home range studies of black rhino have been data deficient and procedurally inconsistent. Standardisation of methods is required.     

Response of native ungulates to drought in semi‐arid Kenyan rangeland

The distribution and abundance of native ungulates were measured on commercially managed, semi‐arid rangeland in central Kenya over a 3‐year period that encompassed severe drought and above‐average rainfall. Native ungulate biomass density averaged 5282 kg km^?2 over the study and was dominated by elephant (Loxodonta africana), impala (Aepyceros melampus) and dik‐dik (Madoqua kirkii). Biomass density of domestic cattle (Bos taurus) averaged 2280 kg km^?2 during the study. Responses of native ungulates to severe drought were variable. Impala densities were similar to or greater than densities for similar habitat in protected areas, and varied from 12 to 16 km^?2 during and following the drought to 24–29 km^?2 following above‐average rainfall. Dik‐dik densities were also greater than densities reported for protected areas and were surprisingly stable throughout the study despite the wide annual fluctuations in rainfall. Elephant migrated out of the region during drought but were present at high densities (2.9–5.2 km^?2) during wet seasons, consistent with telemetry studies emphasizing the importance of Acacia bushland habitat on commercial rangelands for the migratory portion of the Laikipia–Samburu elephant population. Results show that substantial densities of native browsing and mixed‐feeding ungulates can occur on rangeland managed for commercial beef production and suggest that the capacity for ungulates to move over large spatial scales (>100 km²) and to shift distributions in response to locally variable thunderstorms may be important for sustaining these populations.     

Variability in survival and mortality of Acacia drepanolobium Sjøstedt following prescribed burning at Olpejeta Conservancy,Kenya

Mortalities to Acacia drepanolobium, a main item in the diet of the eastern black rhino (Diceros bicornis Michaeli) in Olpejeta conservancy, Kenya, are caused by three main factors: drought, browsers and fire. The effect of controlled fires on A. drepanolobium was examined by monitoring survival and growth in experimental plots before and after fire episodes between 2003 and 2007. Density, survival and growth in A. drepanolobium trees were compared eighteen months before and after burning. Tall trees were more likely to die from fire compared to short trees (R² = 0.575; df = 6; P = 0.048), while seedling densities increased after fire (χ² = 36.57; df = 1; P = 0.001). Even with increased seedling densities, burned areas attracted large numbers of seedling predators, lowering the possibility of seedling recruitment into adult, as mean seedling heights reduced significantly (ANOVA, F = 204.42; df = 1; P = 0.036). Fires also significantly lowered flowering (F = 346; df = 1; P < 0.05) in A. drepanolobium, thereby affecting fruit production. Although fires caused mortalities to adult A. drepanolobium, the most significant effect was tree reversals into seedling height class as trees resprouted. Although fire may increase browse biomass of A. drepanolobium available for black rhino, it is not an appropriate black rhino habitat management tool because burnt areas attract many seedling predators that lower seedling recruitment into adult trees.     

Quantity and Configuration of Available Elephant Habitat and Related Conservation Concerns in the Lower Kinabatangan Floodplain of Sabah,Malaysia

The approximately 300 (298, 95% CI: 152–581) elephants in the Lower Kinabatangan Managed Elephant Range in Sabah, Malaysian Borneo are a priority sub-population for Borneo''s total elephant population (2,040, 95% CI: 1,184–3,652). Habitat loss and human-elephant conflict are recognized as the major threats to Bornean elephant survival. In the Kinabatangan region, human settlements and agricultural development for oil palm drive an intense fragmentation process. Electric fences guard against elephant crop raiding but also remove access to suitable habitat patches. We conducted expert opinion-based least-cost analyses, to model the quantity and configuration of available suitable elephant habitat in the Lower Kinabatangan, and called this the Elephant Habitat Linkage. At 184 km², our estimate of available habitat is 54% smaller than the estimate used in the State''s Elephant Action Plan for the Lower Kinabatangan Managed Elephant Range (400 km²). During high flood levels, available habitat is reduced to only 61 km². As a consequence, short-term elephant densities are likely to surge during floods to 4.83 km⁻² (95% CI: 2.46–9.41), among the highest estimated for forest-dwelling elephants in Asia or Africa. During severe floods, the configuration of remaining elephant habitat and the surge in elephant density may put two villages at elevated risk of human-elephant conflict. Lower Kinabatangan elephants are vulnerable to the natural disturbance regime of the river due to their limited dispersal options. Twenty bottlenecks less than one km wide throughout the Elephant Habitat Linkage, have the potential to further reduce access to suitable habitat. Rebuilding landscape connectivity to isolated habitat patches and to the North Kinabatangan Managed Elephant Range (less than 35 km inland) are conservation priorities that would increase the quantity of available habitat, and may work as a mechanism to allow population release, lower elephant density, reduce human-elephant conflict, and enable genetic mixing.     

A study on the postrelease behaviour and habitat preferences of black rhinos (Diceros bicornis) reintroduced into a fenced reserve in Namibia

Translocations and reintroductions are key elements for the population management of the critically endangered black rhino (Diceros bicornis, Linnaeus, 1758). In this study, we investigated the postrelease behaviour and habitat preferences of a black rhino starter group (n = 4) on the individual level. The animals were reintroduced to a fenced game reserve (87 km²) in North‐Central Namibia 1 year prior to our study. We used camera traps and very high frequency (VHF) radiotelemetry to examine the animals' temporal and spatial behaviour over a period of 4 months at transition between wet and dry seasons. Our results underline a peak in drinking activity and waterhole visits occurring between 7 p.m. and 8 p.m. We found a shift in intensity in drinking activity during the period of the study. Satellite‐based woody cover estimations only suggest positive correlations between the density of woody cover and favoured black rhino habitat types. Although the area seems suitable to facilitate breeding success of this starter group, it does not support a self‐sustaining population. However, black rhinos were already successfully reintroduced to several additional fenced reserves in this region. The selective opening of fences in the future could help to enable genetic exchange between currently isolated groups of rhinos.     

Seasonal diet preferences of black rhinoceros in three arid South African National Parks

Black rhinoceros diet and browse availability was investigated in Augrabies Falls (AFNP), Karoo (KRNP) and Vaalbos National Parks (VNP) in South Africa. Rhino tracks were followed and 18,804 standard bite volumes recorded. Browse availability of each plant species was recorded by measuring 14,800 plants and calculating reachable browse volume. The diet comprised 51, 53 and 41 plant species in AFNP, KRNP and VNP, respectively, but three species accounted for more than 65% of the diet in each park, making diet less diverse than available browse in AFNP and KRNP. Browse availability explained 14%, 15% and 52% of diet selection in AFNP, KRNP and VNP, with consumption of most plant species significantly different from availability. A few plant species were so highly preferred that browsing intensities were unsustainable, while some common species were totally rejected. Pressure on the eaten browse averaged 4.4%, 14.5% and 1.3% annually of the volume in AFNP, KRNP and VNP. In the dry season, there was a two to sevenfold increase in browsing pressure on species with actively photosynthesizing tissues. A few key species, including Monechma and Zygophyllum species, were identified as potential early warning indicators of black rhino browsing impact.     

Woody vegetation damage by the African elephant during severe drought at Pongola Game Reserve,South Africa

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.     

亚洲象生境长时序变化及其对人象冲突的影响——基于融合MaxEnt与HSI模型的生境适宜性评价方法

生境变化直接关系到物种种群维持与人类安全,揭示其长期变化规律及其对人类的影响,可为物种保护与生境恢复提供科学支撑。但因受物种活动点数据获取与位置精度的局限,鲜见濒危、危险物种的长时序生境变化研究。以人象冲突频发的西双版纳勐海—普洱澜沧地区亚洲象种群(勐海—澜沧象群)活动区为例,提出融合MaxEnt与HSI模型的亚洲象长时序生境适宜性评价方法,即基于荟萃分析筛选出的15个亚洲象生境评价因子,结合近期有限的物种活动点监测数据,利用MaxEnt得到生境评价因子的贡献率,再运用HSI模型计算生境适宜性指数;利用该方法制作出研究区1988—2020年逐年时序的亚洲象生境适宜性图,以分析亚洲象生境的时空变化,将其与亚洲象肇事数据结合,进而分析人象冲突与生境变化的关联。结果表明：(1)基于物种生境偏好不变的前提,融合MaxEnt模型与HSI模型的生境适宜性评价方法可应用于物种的长时序生境评价,且基于亚洲象活动点数据从动物对生境利用的生态学视角定量获取亚洲象对各生境评价因子的偏好程度,使生境评价结果具有良好的生态可解释性;(2)目前亚洲象适宜生境面积占研究区面积三分之一(4039.76 km^...     

Dry season habitat selection by a recolonizing population of Asian elephants<Emphasis Type="Italic">Elephas maximus</Emphasis> in lowland Nepal

Owing to landclearing and human expansion, Asian elephantElephas maximus Linnaeus, 1758 is declining throughout its range. In lowland Nepal, the species now only occurs in small remnant populations, shared with India. In order to develop guidelines for conserving the species in the country, we studied the habitat use of a small and recently re-established population in Bardia National Park. We used the distribution of dung in fixed width transects to estimate seasonal habitat selection at a general scale of the Park. We also analyzed a specific habitat selection by elephants within the sal-dominated forest, by comparing the composition of trees and frequency of previous elephant impact on them along fresh tracks with those at random points. Elephants strongly preferred floodplain communities both during the cool and the hot season, but there was a marked shift from forest to grass-dominated subtypes between these seasons. Within the sal-dominated forest, there were more trees with previous elephant impact and a higher density of important food trees, especiallyMallotus phillippinensis along fresh tracks than in random points. We found little if any effect of human activity or location of available water on the spatial distribution of elephant dung. The density of the colonizing population was low (ca 0.2 animals/km²), but numbers are expected to increase in the future. With the preferred floodplain habitat being quite small (ca 60 km²), animals are then expected to spread outside the national park. A large tract of government forest adjacent to the park may then, for some time, provide needed space for the expanding population.     

Updated geographic range maps for giraffe,Giraffa spp., throughout sub‐Saharan Africa,and implications of changing distributions for conservation

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普洱市亚洲象栖息地适宜度评价

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