Scale-dependent selection of greenness by African elephants in the Kruger-private reserve transboundary region,South Africa

Foraging behaviour and habitat selection occur as hierarchical processes. Understanding the factors that govern foraging and habitat selection thus requires investigation of those processes over the scales at which they occur. We investigated patterns of habitat use by African elephants (Loxodonta africana) in relation to vegetation greenness to investigate the scale at which that landscape attribute was most closely related to distribution of elephant locations. We analysed Global Positioning System radio-collar locations for 15 individuals, using the Normalized Difference Vegetation Index as a representation of vegetation greenness in a Geographic Information Systems framework. We compared the importance of vegetation greenness at three spatial scales: the total home range, the seasonal home range and the 16-day home range. During the wet season, seasonal home ranges for both sexes were associated with intermediate greenness within the total home range; there was no evidence of selection based on greenness at finer scales. During the dry season, the strongest associations were within the 16-day home range: individual locations for males tended to be in areas of intermediate greenness, and those for females were in areas of intermediate and high greenness. Our findings suggest that the role of vegetation greenness varies with the scale of analysis, likely reflecting the hierarchical processes involved in habitat selection by elephants.     

Elephant distribution around a volcanic shield dominated by a mosaic of forest and savanna (Marsabit, Kenya)

We investigated the factors that influenced the distribution of the African elephant around a volcanic shield dominated by a mosaic of forest and savanna in northern Kenya. Data on elephant distribution were acquired from four female and five bull elephants, collared with satellite-linked geographical positioning system collars. Based on the eigenvalues (variances) of the correlation matrix, the six factors that contributed significantly to high total variances were distance from drinking water (24%), elevation (15%), shrubland (10%), forest (9%), distance from settlements (8%) and distance from minor roads (7%), contributing to 73% in the observed variation of the elephant distribution. The elephants were found at high forested elevations during the dry season but they moved to the lowlands characterized by shrubland during the wet season. Elevation acts as a proxy for the vegetation structure. The presence of elephants near permanent water points (13%) and seasonal rivers (11%) during the dry and wet seasons, respectively, demonstrates that water is the most important determinant of their distribution throughout the year. We conclude that the distribution of elephants in Marsabit Protected Area and its adjacent areas is influenced mainly by drinking water and vegetation structure.     

Distribution of savannah elephants (Loxodonta africana africana Blumenbach 1797) within Nazinga game ranch,Southern Burkina Faso

We tested the hypothesis that elephant distribution inside the Nazinga Game Ranch (Southern Burkina Faso) during the wet season is influenced by villages outside, while in the dry season elephants are restricted only by water. Occupancy was evaluated by recording elephant dung‐piles on 54 line transects in each of three seasons: wet 2006, dry 2007 and dry 2008. We measured the distance of each transect from nearest villages, nearest permanent water sources, nearest guard posts and tourist camps. The results were unexpected: elephant occupancy in the wet season was independent of villages but influenced by poaching, while their occupancy in both dry seasons was determined by the proximity of villages as well as water. In the dry season, elephants were attracted to villages by grain stores and fruiting trees. There has been a dramatic shift in the dry season distribution of elephants, and consequently in browsing pressure, over the last two decades. We suggest that this change is a consequence of the expansion of human activities outside the ranch. This study shows that the effects of growing human disturbance on elephant populations in small protected areas are not predictable. Correcting for spatial autocorrelation had a negligible effect upon the models.     

A Meta-Analysis of the Impact of African Elephants on Savanna Vegetation

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.     

Relationship between group dynamics and spatial distribution of African elephants in a semi‐arid environment

Group dynamics related to distribution of African elephants in the semi‐arid environment of Tsavo East National Park, Kenya was studied between 2007 and 2010. I examined the seasonal distribution of lone bulls, bull groups, family units and mixed groups and group size. Lone bulls were widely distributed in the dry season and localized in the wet season, whereas bull groups were localized in both seasons. On average bull group size did not differ with seasons or areas, but larger groups were observed in preferred areas in the dry season. Family units were widely distributed in the dry season and localized in the wet season, whereas mixed groups were localized in both seasons. Although family units and mixed groups tended to be large in all areas in the wet season, large groups were also observed in the dry season. Large family units were widely distributed, whereas large mixed groups were localized. The relationship between group dynamics and distribution was attributed to the combination of social, ecological and poaching factors. This study provided insight into distribution patterns that can be applied to the security of the Tsavo elephant population.     

Elephants caught in the middle: impacts of war, fences and people on elephant distribution and abundance in the Caprivi Strip, Namibia

We conducted wet [26 March–4 April 2003 (Apr03)] and dry [1–8 November 2005 (Nov05)] season aerial surveys of African elephants ( Loxodonta africana Blumenbach) in the Caprivi Strip, Namibia to provide an updated status report on elephant numbers and distribution and assist with a historical analysis of elephant distribution and abundance in the Caprivi Strip. During the wet season when water was available in seasonal pans, elephants were widely distributed throughout the survey area. In contrast, during the dry season, a majority of elephant herds occurred within 30 km of the perennial Kwando, Linyanti and Okavango rivers and few herds occurred within the West Caprivi Game Reserve where water in the seasonal pans was limited. We estimated 5318 elephants for the 7731-km² survey area (0.71 elephants km⁻²) for the Apr03 wet season survey and 6474 elephants for the 8597-km² survey area (0.75 elephants km⁻²) for the Nov05 dry season survey. Based on our aerial surveys and reports of elephant numbers and distribution from historical aerial surveys and telemetry studies, civil war, veterinary fences and human activities appear to have effected changes in African elephant abundance, distribution and movements in the Caprivi Strip, Namibia since 1988 when the first comprehensive aerial surveys were conducted.     

Seasonal home ranges of elephants (Loxodonta africana) and their movements between Sabi Sand Reserve and Kruger National Park

We studied the seasonal home ranges and space use of three breeding herds of elephants (Loxodonta africana) for 3–5 years in an area comprising South Africa’s Sabi Sand Reserve (SSR) and Kruger National Park (KNP). Global Positioning System (GPS) transmitters were attached to the matriarchs of three herds and set to transmit a single daily location. Each herd was located in SSR 31%, 60% and 84% of the time. During the wet summer months, the herds walked longer distances and occupied larger seasonal home ranges than during the corresponding dry winter period. Core areas were centred on riverine habitats within both parks, with all three herds exhibiting closer distances to rivers and artificial water holes than would be expected if they were moving randomly. Home ranges within SSR overlapped much of the park. However, in KNP they occupied discrete areas with little overlap. Much of the movement between the two parks occurred along well‐defined corridors. This study shows that elephant herds depended upon the resources of both parks, providing an insight into their within‐ and between‐seasonal movements. This highlights the importance of ongoing co‐operation between wildlife managers from both parks when forming policy.     

Home-range size and movement patterns of Hooded Vultures Necrosyrtes monachus in southern Africa

Hooded Vulture Necrosyrtes monachus populations have declined dramatically in recent years, but we know little about their ecology. We radio-tagged four vultures in northern Botswana to gather data on animal movement and home-range patterns. Hooded Vultures were primarily sedentary at night. Hooded Vultures moved similar distances and speeds during the wet and dry season, and travelled over similar home ranges as measured using minimum convex polygons (MCP), but used much smaller core areas during the dry (breeding) season. We found significant differences in mean distances and speeds moved among different birds, and when comparing day to night, but not between the wet (non-breeding) and dry (breeding) season or by year. All of the variables we tested, including individual vulture differences, season, year and number of fixes, significantly influenced 95% MCP and kernel density estimate (KDE) home-range sizes. Hooded Vultures used significantly smaller KDE home-range sizes during the dry (breeding season) than in the wet (non-breeding) season. Hooded Vultures travelled smaller daily distances over smaller home ranges than most other vulture species for which data exist.     

Ranging behavior of the Asian elephant in Sri Lanka

We studied the ranging patterns of 10 elephants in and around the Yala protected area complex, southern Sri Lanka, using VHF radio telemetry. All tracked elephants displayed similar ranging patterns. The observed home ranges were small (mean=115.2±64.0 km²) relative to reported home ranges in India, possibly in response to high habitat productivity and abundant perennial water sources. Elephants showed high fidelity to their ranges. Home ranges had relatively large core areas, suggesting intensive use of habitat. No geographically distinct seasonal ranges or migratory behavior was observed. Home range overlap was high, and territoriality was absent. Male musth ranges were considerably larger than non-musth ranges and may signify mate searching. Most elephants ranged both in and outside protected areas, suggesting that resources outside protected areas were important for their survival. Thus, translocating and restricting elephants to protected areas will be detrimental to their survival, as it limits resource access. The ranging patterns of Asian elephants suggest that conservation of the species requires their management both in and outside protected areas.     

Understanding the effect of landscape fragmentation and vegetation productivity on elephant habitat utilization in Amboseli ecosystem,Kenya

Understanding factors affecting the distribution of the African elephant is important for its conservation in increasingly human‐dominated savannah landscapes. However, understanding how landscape fragmentation and vegetation productivity affect elephant habitat utilization remains poorly understood. In this study, we tested whether landscape fragmentation and vegetation productivity explain elephant habitat utilization in the Amboseli ecosystem in Kenya. We used GPS (Global Positioning System) telemetry data from five elephants to quantify elephant habitat utilization. Habitat utilization was determined by calculating the time elephants spent within a unit area. We then used generalized additive models (GAMs) to model the relationship between time density and landscape fragmentation, as well as vegetation productivity. Results show that landscape fragmentation and vegetation productivity significantly (P < 0.05) explain elephant habitat utilization. A significant (P < 0.05) unimodal relationship between vegetation productivity and habitat utilization was observed. Results suggest that elephants spend much of their time in less fragmented landscapes of intermediate productivity.     

The ranging patterns of elephants in Marsabit protected area,Kenya: the use of satellite‐linked GPS collars

We investigated the ranging patterns of elephants in the Marsabit protected area, north eastern Kenya, to ascertain the range of bachelor and female family herds in different seasons, and to identify corridor and noncorridor areas. Data were acquired for five bachelor and four female family herds equipped with satellite‐linked geographical positioning system collars, and monitored from December 2005 to December 2007. Distinct dry (about 260 km²) and wet seasons (about 910 km²) ranges were observed, with connecting corridors (north‐eastern corridor: about 90 km long, about 2‐7 km wide; southern corridors: about 10‐20 km long, about 2‐3 km wide). The dry season range corresponded with Marsabit evergreen forest, while the wet season range matched with dry deciduous lowland shrubs. The ranging elephants moved at speed of about 0.2‐20 kmh^?1. Bachelor herds moved faster than female family herds. Elephants moved fast during the intermediate and wet seasons than during the dry season. The speed of ranging elephants was over 1 kmh^?1 in the corridor areas and about 0.2 to less than 1 kmh^?1 in the non‐corridor areas. Expansion of settlements towards corridor areas needs to be controlled to avoid future blocking of connectivity between wet and dry season elephant ranges.     

Soil carbon dynamics and aquatic metabolism of a wet–dry tropics wetland system

Freshwater wetlands are a key component of the global carbon cycle. Wet–dry tropics wetlands function as wet-season carbon sinks and dry-season carbon sources with low aquatic metabolism controlled by predictably seasonal, yet magnitude-variable flow regimes and inundation patterns. However, these dynamics have not been adequately quantified in Australia’s relatively unmodified wet–dry tropics freshwater wetlands. A baseline understanding is required before analysis of land-use or climate change impacts on these aquatic ecosystems can occur. This study characterises geomorphology and sedimentology within a seasonally connected wet–dry tropics freshwater wetland system at Kings Plains, Queensland, Australia, and quantifies soil carbon stocks and wet- and dry-season aquatic metabolism. Soil carbon stocks derived from loss-on-ignition on samples to 1 m depth were 51.5?±?7.8 kg C m^?2, higher than other wet–dry tropics wetlands globally, with potential for long-term retention at greater depths. Gross primary productivity of phytoplankton (GPP) and planktonic respiration (PR) measured through biological oxygen demand bottle experiments in the water column of sediment inundated under laboratory conditions show overall low GPP and PR in both wet- and dry-season samples (all wetland samples were heterotrophic with GPP/PR?<?1). Despite the short-term dominance of aquatic respiration processes leading to net release of carbon in the water column under these conditions, there is appreciable long-term storage of carbon in sediment in the Kings Plains wetlands. This demonstrates the importance of wet–dry-tropics wetland systems as hotspots of carbon sequestration, locally, regionally and globally, and consideration should be given to their conservation and management in this context.

     

Spatial segregation of two vole species (Arvicola sapidus and Microtus cabrerae) within habitat patches in a highly fragmented farmland landscape

Spatial segregation is one of the common mechanisms allowing the co-existence of similar interacting species in heterogeneous environments. Analysing spatial segregation requires information on individual home-range sizes and their degree of spatial overlap. In this study, we used radio-tracking to report for the first time the home-range and core-area sizes of sympatric Cabrera and water voles and to analyse intra- and inter-specific space sharing within habitat patches in a highly fragmented landscape. Results indicated that both species exhibited strong fine-scale site fidelity and reduced variation in range size across sexes and seasons. Monogamous mating system seemed to prevail for both species, although water voles may also exhibit polygynous breeding strategies. Mean home-range and core-area sizes of water voles (946.3 and 156.6 m²) were about twice that of Cabrera voles (418.2 and 55.1 m²). Within habitat patches, individuals of both species often overlapped their home ranges, particularly during the dry season (May–September), though intra-specific home-range overlap was generally higher than inter-specific overlap. Inter-specific space sharing was restricted to areas outside the centre of activity of animals, as no core-area overlap was ever recorded between Cabrera and water voles. Taken together, results support the view that co-existence of Cabrera and water voles in Mediterranean patchy habitats may in part result from spatial segregation among individuals, which may reflect competitive displacement or small-scale habitat partitioning. Results highlight the need to account for species interactions when designing conservation management strategies for sympatric Cabrera and water voles in fragmented landscapes.     

Spatial distribution and movements of elephants (Loxodonta africana) in the northern Namib Desert region of the Kaokoveld, South West Africa/Namibia

The home ranges, space-use patterns, seasonal distribution, long-distance movements, water-related movements and movement patterns of elephants ( Loxodonta africana ) residing in the northern Namib Desert region of the Kaokoveld, South West Africa/Namibia, are discussed. All movements of the desert-dwelling elephants, including occasional wanderings up to 195 km, were confined to the northern Namib Desert and no evidence could be found of any movements including migrations to and from the Etosha National Park. Home range size varied from 1763 to 2944 km² and the main feature of the spatial distribution of these elephants is the marked fidelity to individual home ranges. Seasonal distribution and space-use patterns within individual home ranges coincide with seasonal climatic changes and the corresponding changes in food and water availability. The elephant's normal movements and migrations were confined to individual home ranges and no mass population migrations took place. Their mobility and their ability to go up to four days without drinking water enabled them to utilize food sources up to 70 km away from waterholes and, together with their intimate knowledge about resource distribution within their home ranges, are regarded as key factors in their survival in the desert. The elephants' spatial distribution and movement patterns indicate a natural population segregation from other elephant populations in the Kaokoveld, clearly demarcating the desert-dwelling elephants as a distinct population with their own movement patterns and geographic range.     

Elephant (Loxodonta africana) Home Ranges in Sabi Sand Reserve and Kruger National Park: A Five-Year Satellite Tracking Study

During a five-year GPS satellite tracking study in Sabi Sand Reserve (SSR) and Kruger National Park (KNP) we monitored the daily movements of an elephant cow (Loxodonta africana) from September 2003 to August 2008. The study animal was confirmed to be part of a group of seven elephants therefore her position is representative of the matriarchal group. We found that the study animal did not use habitat randomly and confirmed strong seasonal fidelity to its summer and winter five-year home ranges. The cow''s summer home range was in KNP in an area more than four times that of her SSR winter home range. She exhibited clear park habitation with up to three visits per year travelling via a well-defined northern or southern corridor. There was a positive correlation between the daily distance the elephant walked and minimum daily temperature and the elephant was significantly closer to rivers and artificial waterholes than would be expected if it were moving randomly in KNP and SSR. Transect lines established through the home ranges were surveyed to further understand the fine scale of the landscape and vegetation representative of the home ranges.     

Home-Range Attributes of Raccoons in a Fragmented Agricultural Region of Northern Indiana

Abstract: For many wildlife species, agricultural landscapes undergo spatial and temporal fluctuations in the composition of food and cover annually with the planting and harvesting of crops. Raccoon (Procyon lotor) populations have flourished in agricultural landscapes, where crops increase foraging opportunities and efficiencies. However, information is lacking regarding the effects of temporal shifts in food and cover resulting from agricultural activities on raccoon home ranges. We examined home-range characteristics of 60 (33 M, 27 F) adult raccoons in northern Indiana, USA, from May 2003 through June 2005 to identify shifts in the size of home ranges and core use areas among seasons defined by crop availability and crop developmental stages. Mean fixed-kernel home-range (92 ± 6 ha; xM ± SE) and core-area sizes (20 ± 2 ha) of males were significantly larger than those of females (58 ± 7 ha and 13 ± 2 ha, respectively), and both were smaller than those reported for raccoons in other fragmented agricultural landscapes. Home-range sizes varied little among seasons for either sex. However, home ranges of males were smallest during the crop maturation stage, whereas home ranges of females were smallest during the crop growing season. The results of our study suggest that even in expansive rural landscapes, raccoons can maintain small home ranges when food, water, and denning resources are readily available. Additionally, the lack of differences among seasonal home-range sizes, despite the presence of an ephemeral superabundant food source (i.e., corn) during the maturation season, was likely due to the close proximity of foraging and denning resources across seasons.     

Linking Vital Rates of Landbirds on a Tropical Island to Rainfall and Vegetation Greenness

Remote tropical oceanic islands are of high conservation priority, and they are exemplified by range-restricted species with small global populations. Spatial and temporal patterns in rainfall and plant productivity may be important in driving dynamics of these species. Yet, little is known about environmental influences on population dynamics for most islands and species. Here we leveraged avian capture-recapture, rainfall, and remote-sensed habitat data (enhanced vegetation index [EVI]) to assess relationships between rainfall, vegetation greenness, and demographic rates (productivity, adult apparent survival) of three native bird species on Saipan, Northern Mariana Islands: rufous fantail (Rhipidura rufifrons), bridled white-eye (Zosterops conspicillatus), and golden white-eye (Cleptornis marchei). Rainfall was positively related to vegetation greenness at all but the highest rainfall levels. Temporal variation in greenness affected the productivity of each bird species in unique ways. Predicted productivity of rufous fantail was highest when dry and wet season greenness values were high relative to site-specific 5-year seasonal mean values (i.e., relative greenness); while the white-eye species had highest predicted productivity when relative greenness contrasted between wet and dry seasons. Survival of rufous fantail and bridled white eye was positively related to relative dry-season greenness and negatively related to relative wet-season greenness. Bridled white-eye survival also showed evidence of a positive response to overall greenness. Our results highlight the potentially important role of rainfall regimes in affecting population dynamics of species on oceanic tropical islands. Understanding linkages between rainfall, vegetation, and animal population dynamics will be critical for developing effective conservation strategies in this and other regions where the seasonal timing, extent, and variability of rainfall is expected to change in the coming decades.     

Response of African Elephants (Loxodonta africana) to Seasonal Changes in Rainfall

The factors that trigger sudden, seasonal movements of elephants are uncertain. We hypothesized that savannah elephant movements at the end of the dry season may be a response to their detection of distant thunderstorms. Nine elephants carrying Global Positioning System (GPS) receivers were tracked over seven years in the extremely dry and rugged region of northwestern Namibia. The transition date from dry to wet season conditions was determined annually from surface- and satellite-derived rainfall. The distance, location, and timing of rain events relative to the elephants were determined using the Tropical Rainfall Measurement Mission (TRMM) satellite precipitation observations. Behavioral Change Point Analysis (BCPA) was applied to four of these seven years demonstrating a response in movement of these elephants to intra- and inter-seasonal occurrences of rainfall. Statistically significant changes in movement were found prior to or near the time of onset of the wet season and before the occurrence of wet episodes within the dry season, although the characteristics of the movement changes are not consistent between elephants and years. Elephants in overlapping ranges, but following separate tracks, exhibited statistically valid non-random near-simultaneous changes in movements when rainfall was occurring more than 100 km from their location. While the environmental trigger that causes these excursions remains uncertain, rain-system generated infrasound, which can travel such distances and be detected by elephants, is a possible trigger for such changes in movement.     

Landscape-scale habitat response of African elephants shows strong selection for foraging opportunities in a human dominated ecosystem

Balancing trade-offs between foraging and risk factors is a fundamental behavior that structures the spatial distribution of species. For African elephants Loxodonta africana, human pressures from poaching and conflict are primary drivers of species decline, but little is known about how elephants structure their spatial behavior in the face of human occupancy and predation. We seek to understand how elephants balance trade-offs between resource access, human presence and human predatory risk factors (poaching and conflict killing) in an unfenced, dynamic ecosystem where elephants persist primarily outside protected areas in community rangelands. We used tracking data from 101 elephants collected between 2001 and 2016. We investigated elephant behavior in response to landcover, topography, productivity, water, human features and human predation risk using third-order resource selection functions. We extended this analysis by employing a mixed-effects multinomial regression to identify temporal shifts in habitat use, and evaluated temporal shifts in movement patterns by estimating mean squared displacement across different productivity periods. Across periods, elephants displayed strong selection for productive areas and areas near water. Temporal shifts in habitat use showed that, during the dry period, elephants were clustered around permanent water sources where humans also congregated. At the onset of the wet period, a shift occurred where elephants moved away from permanent water and from permanent settlements towards seasonal water sources and seasonal settlements. Our findings indicate that foraging and water access are important limiting factors affecting elephants that potentially restrain their spatial responses to humans at the scale of our analysis. Given that pastoralists and elephants rely on the same resources, increasing human and livestock populations enhance pressure on shared resources and space in Africa's drylands. The long-term conservation of elephants will require approaches that reduce poaching as well as landscape level planning to prevent negative impacts from increasing competition for preferred resources.     

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.)