Precipitation, vegetation productivity, and human impacts control home range size of elephants in dryland systems in northern Namibia.

Climatic variability, resource availability, and anthropogenic impacts heavily influence an animal's home range. This makes home range size an effective metric for understanding how variation in environmental factors alter the behavior and spatial distribution of animals. In this study, we estimated home range size of African elephants (Loxodonta africana) across four sites in Namibia, along a gradient of precipitation and human impact, and investigated how these gradients influence the home range size on regional and site scales. Additionally, we estimated the time individuals spent within protected area boundaries. The mean 50% autocorrelated kernel density estimate for home range was 2200 km2 [95% CI:1500-3100 km2]. Regionally, precipitation and vegetation were the strongest predictors of home range size, accounting for a combined 53% of observed variation. However, different environmental covariates explained home range variation at each site. Precipitation predicted most variation (up to 74%) in home range sizes (n = 66) in the drier western sites, while human impacts explained 71% of the variation in home range sizes (n = 10) in Namibia's portion of the Kavango-Zambezi Transfrontier Conservation Area. Elephants in all study areas maintained high fidelity to protected areas, spending an average of 85% of time tracked on protected lands. These results suggest that while most elephant space use in Namibia is driven by natural dynamics, some elephants are experiencing changes in space use due to human modification.

THE PULMONARY AND METABOLIC EFFECTS OF SUSPENSION BY THE FEET COMPARED WITH LATERAL RECUMBENCY IN IMMOBILIZED BLACK RHINOCEROSES (DICEROS BICORNIS) CAPTURED BY AERIAL DARTING.

Aerial translocation of captured black rhinoceroses (Diceros bicornis) has been accomplished by suspending them by their feet. We expected this posture would compromise respiratory gas exchange more than would lateral recumbency. Because white rhinoceroses (Ceratotherium simum) immobilized with etorphine alone are hypermetabolic, with a high rate of carbon dioxide production (VCO2), we expected immobilized black rhinoceroses would also have a high VCO2. Twelve (nine male, three female; median age 8 yr old [range: 4-25]; median weight 1,137 kg [range: 804-1,234] body weight) wild black rhinoceroses were immobilized by aerial darting with etorphine and azaperone. The animals were in lateral recumbency or suspended by their feet from a crane for approximately 10 min before data were collected. Each rhinoceros received both treatments sequentially, in random order. Six were in lateral recumbency first and six were suspended first. All animals were substantially hypoxemic and hypercapnic in both postures. When suspended by the feet, mean arterial oxygen pressure (PaO2) was 42 mm Hg, 4 mm Hg greater than in lateral recumbency (P=0.030), and arterial carbon dioxide pressure (PaCO2) was 52 mm Hg, 3 mm Hg less than in lateral recumbency (P=0.016). Tidal volume and minute ventilation were similar between postures. The mean VCO2 was 2 mL/kg/min in both postures and was similar to, or marginally greater than, VCO2 predicted allometrically. Suspension by the feet for 10 min did not impair pulmonary function more than did lateral recumbency and apparently augmented gas exchange to a small degree relative to lateral recumbency. The biological importance in these animals of numerically small increments in PaO2 and decrements in PaCO2 with suspension by the feet is unknown. Black rhinoceroses immobilized with etorphine and azaperone were not as hypermetabolic as were white rhinoceroses immobilized with etorphine.

Mapping and assessing the impact of small-scale ephemeral water sources on wildlife in an African seasonal savannah.

In many savannah regions of Africa, pronounced seasonal variability in rainfall results in wildlife being restricted to floodplains and other habitats adjacent to permanent surface water in the dry season. During the wet season, rainfall fills small-scale, ephemeral water sources that allow wildlife to exploit forage and other resources far from permanent surface water. These water sources remain difficult to quantify, however, due to their small and ephemeral nature, and as a result are rarely included in quantitative studies of wildlife distribution, abundance, and movement. Our goal was to map ephemeral water in Bwabwata National Park in Namibia using two different approaches and to relate measures of ephemeral water to the abundance, distribution, and movement of two large wildlife species. We used high-resolution Google Earth and Esri World imagery to visually identify waterholes. Additionally, we used Sentinel-2 satellite imagery to map ephemeral water across the study area using the Normalized Difference Water Index. With these mapped waterhole layers and data from GPS-collared individuals of African elephant (Loxodonta africana) and African buffalo (Syncerus caffer), we evaluated the importance of ephemeral water in conditioning abundance and movement of these two species. The two approaches to mapping ephemeral water resulted in the visual identification of nearly 10,000 waterholes, and a predicted ephemeral water layer of ~76% accuracy. The inclusion of ephemeral water into models of abundance and movement resulted in improved goodness of fit relative to those without water, and water impacts on abundance and movement were among the strongest of all variables considered. The potential importance of ephemeral water in conditioning the movements and distributions of large herbivores in African savannahs has been difficult to quantify relative to vegetation drivers. Our results suggest research into ephemeral water impacts deserves more attention.

Phylogeography, genetic diversity, and population structure of Nile crocodile populations at the fringes of the southern African distribution.

Nile crocodiles are apex predators widely distributed in sub-Saharan Africa that have been viewed and managed as a single species. A complex picture of broad and fine-scale phylogeographic patterns that includes the recognition of two species (Crocodylus niloticus and Crocodylus suchus), and the structuring of populations according to river basins has started to emerge. However, previous studies surveyed a limited number of samples and geographical regions, and large areas of the continent remained unstudied. This work aimed at a fine scale portrait of Nile crocodile populations at the fringes of their geographic distribution in southern Africa. Wild and captive individuals were sampled across four major river systems (Okavango, Lower Kunene, Lower Shire and Limpopo) and the KwaZulu-Natal region. A multi-marker approach was used to infer phylogeographic and genetic diversity patterns, including new and public mitochondrial data, and a panel of 11 nuclear microsatellites. All individuals belonged to a phylogenetic clade previously associated with the C. niloticus species, thus suggesting the absence of C. suchus in southern Africa. The distribution of mitochondrial haplotypes indicated ancestral genetic connectivity across large areas, with loss of diversity along the north-south axis. Genetic variation partitioned the populations primarily into western and eastern regions of southern Africa, and secondarily into the major river systems. Populations were partitioned into five main groups corresponding to the Lower Kunene, the Okavango, the Lower Shire, and the Limpopo rivers, and the KwaZulu-Natal coastal region. All groups show evidence of recent bottlenecks and small effective population sizes. Long-term genetic diversity is likely to be compromised, raising conservation concern. These results emphasize the need for local genetic assessment of wild populations of Nile crocodiles to inform strategies for management of the species in southern Africa.

Effects of age and sex ratios on offspring recruitment rates in translocated black rhinoceros.

Success of animal translocations depends on improving postrelease demographic rates toward establishment and subsequent growth of released populations. Short-term metrics for evaluating translocation success and its drivers, like postrelease survival and fecundity, are unlikely to represent longer-term outcomes. We used information theory to investigate 25 years of data on black rhinoceros (Diceros bicornis) translocations. We used the offspring recruitment rate (ORR) of translocated females-a metric integrating survival, fecundity, and offspring recruitment at sexual maturity-to detect determinants of success. Our unambiguously best model (AICω = 0.986) predicted that ORR increases with female age at release as a function of lower postrelease adult rhinoceros sex ratio (males:females). Delay of first postrelease reproduction and failure of some females to recruit any calves to sexual maturity most influenced the pattern of ORRs, and the leading causes of recruitment failure were postrelease female death (23% of all females) and failure to calve (24% of surviving females). We recommend translocating older females (≥6 years old) because they do not exhibit the reproductive delay and low ORRs of juveniles (<4 years old) or the higher rates of recruitment failure of juveniles and young adults (4-5.9 years old). Where translocation of juveniles is necessary, they should be released into female-biased populations, where they have higher ORRs. Our study offers the unique advantage of a long-term analysis across a large number of replicate populations-a science-by-management experiment as a proxy for a manipulative experiment, and a rare opportunity, particularly for a large, critically endangered taxon such as the black rhinoceros. Our findings differ from previous recommendations, reinforce the importance of long-term data sets and comprehensive metrics of translocation success, and suggest attention be shifted from ecological to social constraints on population growth and species recovery, particularly when translocating species with polygynous breeding systems.

Experimental evidence for homeostatic sex allocation after sex-biased reintroductions.

First principles predict negative frequency-dependent sex allocation, but it is unproven in field studies and seldom considered, despite far-reaching consequences for theory and practice in population genetics and dynamics as well as animal ecology and behaviour. Twenty-four years of rhinoceros calving after 45 reintroductions across southern Africa provide the first in situ experimental evidence that unbalanced operational sex ratios predicted offspring sex and offspring sex ratios. Our understanding of population dynamics, especially reintroduction and invasion biology, will be significantly impacted by these findings.

Pulmonary dead space in free-ranging immobilized black rhinoceroses (Diceros bicornis) in Namibia.

It was observed previously that end-expired carbon dioxide (P(E)CO2) decreased when immobilized black rhinoceroses (Diceros bicornis) were moved from sternal to lateral recumbency. These experiments were designed to test whether greater alveolar ventilation or greater pulmonary dead space in lateral recumbency explains this postural difference in P(E)CO2. Twenty-one (9 male, 12 female; 15 [3.5-26] yr old) wild black rhinoceroses were immobilized with etorphine and azaperone and positioned in either sternal or lateral recumbency. All rhinoceroses were hypoxemic and had lactic and respiratory acidemia. The animals in lateral recumbency were more acidemic, had higher lactate, and lower arterial oxygen that those in sternal recumbency; however, arterial carbon dioxide was similar between groups. Both P(E)CO2 and mixed expired carbon dioxide pressure were lower in lateral than sternal recumbency. Although there was no difference in tidal volume or arterial carbon dioxide, both the breathing rate and minute ventilation were greater in lateral recumbency. The physiologic dead space ratio and dead space volume were approximately two times larger in lateral recumbency; hence, the decrease in P(E)CO2 in lateral recumbency can be attributed to increased dead space ventilation not increased alveolar ventilation. Positioning immobilized rhinoceroses in lateral recumbency does not confer any advantage over sternal in terms of ventilation, and the increase in minute ventilation in lateral recumbency can be considered an energetic waste. Although arterial oxygen was superior in sternal recumbency, further studies that measure oxygen delivery (e.g., to the muscles of locomotion) are warranted before advice regarding the optimal position for immobilized rhinoceroses can be given with confidence.

Contrasting historical and recent gene flow among African buffalo herds in the Caprivi Strip of Namibia.

Population genetic structure is often used to infer population connectivity, but genetic structure may largely reflect historical rather than recent processes. We contrasted genetic structure with recent gene-flow estimates among 6 herds of African buffalo (Syncerus caffer) in the Caprivi Strip, Namibia, using 134 individuals genotyped at 10 microsatellite loci. We tested whether historical and recent gene flows were influenced by distance, potential barriers (rivers), or landscape resistance (distance from water). We also tested at what scales individuals were more related than expected by chance. Genetic structure across the Caprivi Strip was weak, indicating that historically, gene flow was strong and was not affected by distance, barriers, or landscape resistance. Our analysis of simulated data suggested that genetic structure would be unlikely to reflect human disturbances in the last 10-20 generations (75-150 years) because of slow predicted rates of genetic drift, but recent gene-flow estimates would be affected. Recent gene-flow estimates were not consistently affected by rivers or distance to water but showed that isolation by distance appears to be developing. Average relatedness estimates among individuals exceeded random expectations only within herds. We conclude that historically, African buffalo moved freely throughout the Caprivi Strip, whereas recent gene flow has been more restricted. Our findings support efforts to maintain the connectivity of buffalo herds across this region and demonstrate the utility of contrasting genetic inferences from different time scales.

Home on the range: factors explaining partial migration of African buffalo in a tropical environment.

Partial migration (when only some individuals in a population undertake seasonal migrations) is common in many species and geographical contexts. Despite the development of modern statistical methods for analyzing partial migration, there have been no studies on what influences partial migration in tropical environments. We present research on factors affecting partial migration in African buffalo (Syncerus caffer) in northeastern Namibia. Our dataset is derived from 32 satellite tracking collars, spans 4 years and contains over 35,000 locations. We used remotely sensed data to quantify various factors that buffalo experience in the dry season when making decisions on whether and how far to migrate, including potential man-made and natural barriers, as well as spatial and temporal heterogeneity in environmental conditions. Using an information-theoretic, non-linear regression approach, our analyses showed that buffalo in this area can be divided into 4 migratory classes: migrants, non-migrants, dispersers, and a new class that we call "expanders". Multimodel inference from least-squares regressions of wet season movements showed that environmental conditions (rainfall, fires, woodland cover, vegetation biomass), distance to the nearest barrier (river, fence, cultivated area) and social factors (age, size of herd at capture) were all important in explaining variation in migratory behaviour. The relative contributions of these variables to partial migration have not previously been assessed for ungulates in the tropics. Understanding the factors driving migratory decisions of wildlife will lead to better-informed conservation and land-use decisions in this area.

Serial temperature monitoring and comparison of rectal and muscle temperatures in immobilized free-ranging black rhinoceros (Diceros bicornis).

Control of body temperature is critical to a successful anesthetic outcome, particularly during field immobilization of wild animals. Hyperthermia associated with exertion can lead to serious and potentially life-threatening complications such as organ damage (including myopathy) and death. Methods for monitoring core body temperature must accurately reflect the physiologic status of the animal in order for interventions to be effective. The goal of this preliminary study was to compare serial rectal and muscle temperatures in field-immobilized black rhinoceros (Diceros bicornis) and evaluate a possible association. Twenty-four free-ranging black rhinoceros were immobilized between February and March of 2010 in Ethosha National Park, Namibia. Pairwise comparisons showed a correlation of 0.73 (95% CI; 0.70-0.75) between rectal and muscle temperature measurements. Results from a multivariable model indicate that muscle temperature readings were, on average, 0.46 degrees C (95% CI; 0.36-0.57 degrees C) higher than rectal temperatures while adjusting for repeated measurements on the same rhinoceros, effect of duration of immobilization, and effect of ambient temperature on rhinoceroses' temperature readings. As immobilization time increased, muscle and rectal temperature values within an individual rhinoceros tended to equilibrate. The overall temperatures decreased by an average of 0.00059 degrees C/min (95% CI; -0.0047 to -0.0035 degrees C/min; P = 0.779). As the ambient temperature at time of immobilization increased by 1 degree C, the average rhinoceros temperature increased by 0.09 degrees C (95% CI; 0.06-0.11 degrees C, P < 0.0001). Higher body temperature creates a potential for cellular damage leading to complications that include myopathy. Methods for monitoring rectal, muscle, and ambient temperatures should be incorporated into anesthetic monitoring protocols for large ungulates, particularly under field conditions.

Translocations as experiments in the ecological resilience of an asocial mega-herbivore.

Species translocations are remarkable experiments in evolutionary ecology, and increasingly critical to biodiversity conservation. Elaborate socio-ecological hypotheses for translocation success, based on theoretical fitness relationships, are untested and lead to complex uncertainty rather than parsimonious solutions. We used an extraordinary 89 reintroduction and 102 restocking events releasing 682 black rhinoceros (Diceros bicornis) to 81 reserves in southern Africa (1981-2005) to test the influence of interacting socio-ecological and individual characters on post-release survival. We predicted that the socio-ecological context should feature more prominently after restocking than reintroduction because released rhinoceros interact with resident conspecifics. Instead, an interaction between release cohort size and habitat quality explained reintroduction success but only individuals' ages explained restocking outcomes. Achieving translocation success for many species may not be as complicated as theory suggests. Black rhino, and similarly asocial generalist herbivores without substantial predators, are likely to be resilient to ecological challenges and robust candidates for crisis management in a changing world.