Etosha National Park Research Articles

Population genomics of the southern giraffe

Studying wildlife taxonomic diversity and identifying distinct populations has traditionally been largely based on morphology and geographic origin. More recently, this method has been supplemented by genetic data from the mitochondrial genome. However, this is limited as only maternally inherited and may not reflect the true nature of a population’s genetics. Within the giraffe (Giraffa spp.), subspecies and unique populations were successfully characterized using both mitochondrial and genomic DNA studies, which led to new insights and, in some cases, unexpected results that required further verification. Here, we sequenced the genomes of 85 southern giraffe (G. giraffa) individuals from ten populations across southern Africa for a detailed investigation into the genetic diversity and history of its two subspecies, the Angolan (G. g. angolensis) and the South African (G. g. giraffa) giraffe. While the overall genotypes show low levels of runs of homozygosity compared to other mammals, the degree of heterozygosity is limited despite the large population size of South African giraffe. The nuclear genotype is largely congruent with the mitochondrial genotype. However, we have identified that the distribution of the Angolan giraffe is not as far east as indicated in an earlier mitochondrial DNA study. Botswana’s Central Kalahari Game Reserve giraffe are unique, with a clear admixture of Angolan and South African giraffe populations. However, the enigmatic desert-dwelling giraffe of northwest Namibia is locally distinct from other Angolan giraffe yet exhibits intra-subspecies signs of admixture resulting from a recent introduction of individuals from Namibia’s Etosha National Park. Whole genome sequencing is an invaluable and nearly indispensable tool for wildlife management to uncover genetic diversity that is undetectable through mitogenomic, geographical, and morphological means.

Identification of landscape features structuring movement connectivity for Namibian elephants

ContextHuman modification of landscapes poses a significant threat to wildlife, particularly in Africa where population growth and land conversion are expected to increase. Habitat loss and fragmentation have led to declines in wildlife populations, highlighting the need to identify and preserve critical habitats, including core use areas and connectivity between them. Most recently, the identification of habitat corridors has become a key objective.ObjectivesOur study objectives are to (1) empirically quantify connectivity across the Kunene—Etosha landscape in Northwestern Namibia using GPS tracking data on wild African elephants, and (2) assess the landscape features (i.e., geologic, biotic, and human-made) influencing connectivity and corridor types (e.g., fast movement corridors versus slow multi-use movement corridors).MethodsWe used GPS telemetry data from 66 elephants collared in Northwestern Namibia to empirically quantify connectivity using a graph theoretic approach and assess landscape features influencing connectivity. Based on the ‘movescape’ approach, we identify different types of corridors and examined how landscape features differed across these corridors using multiple regression models on locations classified into different types of use categories by machine learning algorithms.ResultsOur results revealed strong variation in connectivity across the landscape, with paths of high connectivity near water sources between the study areas. We found that factors related to water sources and human presence primarily influenced connectivity. Water holes serve as hubs across the ecosystem for both male and female elephants with lower use areas peripheral to areas with water. Connectivity between Kunene and Etosha National Park was relatively rare among the collared elephants, but we highlight the key areas used to move between the two regions.ConclusionWater was the key feature structuring space use, and human presence influenced connectivity between water points, highlighting the importance of landscape planning in relation to limited water sources and human activities. Our results suggest that focusing management efforts on areas where water is limited for both elephants and humans will be important to reduce conflict and maintain ecosystem connectivity.

Crossing Etosha: a history of donkeys in Namibia’s central north

Donkeys are a common sight in Namibia’s central north and play an important albeit contested role in rural society and economy. Historically, they were one of the few domesticated animals that were allowed to enter today’s Etosha National Park and to cross the veterinary border (Red Line) between the northern communal lands and the Police Zone in central Namibia. In the first half of the twentieth century, migrant workers brought thousands of donkeys from the south to what was then called “Ovamboland.” Even after the import was regulated more strictly in the 1940s, donkeys could still get permission to be used as transport animals to carry goods through Etosha. We retrace the routes of the domesticated donkey through a conservation landscape of wildlife and explore entanglements between donkeys, other livestock and wildlife in and out of Etosha. We ask how the particular legal position of the donkey — and its close links to migrant labour — impacted multispecies entanglements in past rural societies and how this legal position has changed over time, including in independent Namibia. We provide a nuanced understanding of the contested position of the donkey in northern Namibia, using examples from King Nehale Conservancy north of Etosha.

The use of vocal coordination in male African elephant group departures: evidence of active leadership and consensus.

Group-living animals engage in coordinated vocalizations to depart from a location as a group, and often, to come to a consensus about the direction of movement. Here, we document for the first time, the use of coordinated vocalizations, the "let's go" rumble, in wild male African elephant group departures from a waterhole. We recorded vocalizations and collected behavioral data as known individuals engaged in these vocal bouts during June-July field seasons in 2005, 2007, 2011, and 2017 at Mushara waterhole within Etosha National Park, Namibia. During departure events, we documented which individuals were involved in the calls, the signature structure of each individual's calls, as well as the ordering of callers, the social status of the callers, and those who initiated departure. The "let's go" rumble was previously described in tight-knit family groups to keep the family together during coordinated departures. Male elephants are described as living in loose social groups, making this finding particularly striking. We found that this vocal coordination occurs in groups of closely associated, highly bonded individuals and rarely occurs between looser associates. The three individuals most likely to initiate the "let's go" rumble bouts were all highly socially integrated, and one of these individuals was also the most dominant overall. This finding suggests that more socially integrated individuals might be more likely to initiate, or lead, a close group of associates in the context of leaving the waterhole, just as a high-ranking female would do in a family group. The fact that many individuals were involved in the vocal bouts, and that departure periods could be shorter, longer, or the same amount of time as pre-departure periods, all suggest that there is consensus with regard to the act of leaving, even though the event was triggered by a lead individual.

Effect of the habitat and tusks on trunk grasping techniques in African savannah elephants.

Among tetrapods, grasping is an essential function involved in many vital behaviours. The selective pressures that led to this function were widely investigated in species with prehensile hands and feet. Previous studies namely highlighted a strong effect of item properties but also of the species habitat on manual grasping behaviour. African savannah elephants (Loxodonta africana) are known to display various prehensile abilities and use their trunk in a large diversity of habitats. Composed of muscles and without a rigid structure, the trunk is a muscular hydrostat with great freedom of movement. This multitasking organ is particularly recruited for grasping food items while foraging. Yet, the diet of African savannah elephants varies widely between groups living in different habitats. Moreover, they have tusks alongside the trunk which can assist in grasping behaviours, and their tusk morphologies are known to vary considerably between groups. Therefore, in this study, we investigate the food grasping techniques used by the trunk of two elephant groups that live in different habitats: an arid study site in Etosha National Park in Namibia, and an area with consistent water presence in Kruger National Park in South Africa. We characterised the tusks profiles and compared the grasping techniques and their frequencies of use for different foods. Our results show differences in food-grasping techniques between the two groups. These differences are related to the food item property and tusk profile discrepancies highlighted between the two groups. We suggest that habitat heterogeneity, particularly aridity gaps, may induce these differences. This may reveal an optimisation of grasping types depending on habitat, food size and accessibility, as well as tusk profiles.

Spotted hyena (Crocuta crocuta) predation on passerine birds in Namibia

Spotted hyenas (Crocuta crocuta) are known for their flexible foraging behavior and trophic interactions with diverse prey species. However, most studies describing their hunting behavior are focused on predation of large prey. Here we describe the capture rates and behavior of adult and subadult spotted hyenas hunting passerine birds. Hyenas were actively chasing, catching and feeding on red-billed queleas (Quelea quelea), a passerine bird gathering in large flocks at a waterhole in the Etosha National Park, Namibia. In total, we observed 38 successful captures with the average individual capture rate of 21 birds caught per hour. It remains unclear if this is a learned behavior of a single hyena clan or if it could be a widespread interaction across Africa, where the two species co-occur. Although it is unlikely that passerines could make up a substantial part of a spotted hyenas diet regardless of the location, our observations provide a rare documented example of trophic interactions between a large carnivore and small birds. It also represents an addition to a diverse repertoire of hyena foraging behaviors, which confirms their adaptability in obtaining food from non-typical sources.

Season of death, pathogen persistence and wildlife behaviour alter number of anthrax secondary infections from environmental reservoirs.

An important part of infectious disease management is predicting factors that influence disease outbreaks, such as R, the number of secondary infections arising from an infected individual. Estimating R is particularly challenging for environmentally transmitted pathogens given time lags between cases and subsequent infections. Here, we calculated R for Bacillus anthracis infections arising from anthrax carcass sites in Etosha National Park, Namibia. Combining host behavioural data, pathogen concentrations and simulation models, we show that R is spatially and temporally variable, driven by spore concentrations at death, host visitation rates and early preference for foraging at infectious sites. While spores were detected up to a decade after death, most secondary infections occurred within 2 years. Transmission simulations under scenarios combining site infectiousness and host exposure risk under different environmental conditions led to dramatically different outbreak dynamics, from pathogen extinction (R < 1) to explosive outbreaks (R > 10). These transmission heterogeneities may explain variation in anthrax outbreak dynamics observed globally, and more generally, the critical importance of environmental variation underlying host-pathogen interactions. Notably, our approach allowed us to estimate the lethal dose of a highly virulent pathogen non-invasively from observational studies and epidemiological data, useful when experiments on wildlife are undesirable or impractical.

Impact of Ungulates on Vegetation Composition Around Waterholes in the Western Part of Etosha National Park, Namibia

Etosha National Park (ENP) is one of the largest conservation areas located in the semi-arid regions of Namibia. Artificial waterholes (boreholes) are the main sources of water for game in the park. Water availability has been largely viewed as a major factor driving ungulate’s impact on vegetation composition around waterholes. A nested-intensity sampling design was adopted to collect data from fifty-four (54) quadrats (25m x 25m) on six transects (two at each waterhole) measuring 1 800m from Renostervlei, Dolomietpunt and Olifantsrus waterholes. Results showed that there was no significant difference in species composition with increasing distance away from the waterholes. Herbivory, soil trampling, playing and fighting of ungulates among vegetation before and after drinking has impacted vegetation composition around waterholes. The impacts imposed on vegetation around waterholes by ungulates cannot be completely avoided but be reduced. To reduce the ungulate’s impacts on vegetation composition around waterholes, adaptive management measures such as controlled burning, closure of waterholes during rainy seasons, creation of new waterholes that will be evenly distributed within the park and destocking of ungulates is recommended to restore some of the plant species that may have been lost from within those localities.

A Snapshot into the Lives of Elephants: Camera Traps and Conservation in Etosha National Park, Namibia

Knowledge of elephant movement and grouping patterns in the wild is critical for their management and conservation. Much of these data come from GPS collar data and aerial surveys, which have provided invaluable information, but data from these methods are often limited to small groups or entire populations. Effective elephant management requires both generalized and localized methodologies. Here, we propose the expanded use of camera traps in research relating to elephant localized movements and grouping patterns as an additional tool for elephant conservation management. In this study, we use a battery-powered camera trap to provide daily high-resolution data of African savanna elephant (Loxodonta africana) grouping patterns over the course of an entire year. We present findings on the seasonal and diurnal grouping patterns of elephants at a waterhole in the northeast corner of Etosha National Park from July 2016 to June 2017. The frequency of elephant occurrences varied seasonally and diurnally across all group types (solitary male, male, family, and mixed groups), while group sizes did not vary seasonally, except for male groups. Solitary males occurred relatively equally throughout the day, while male and mixed groups occurred the most midday, and family groups occurred the most in the afternoon. Additionally, we measured the reliability of research assistants when collecting group type and group size data from the camera trap images. Intra- and inter-observer reliability was excellent among and across research assistants, highlighting the potential for non-specialist observers to have greater involvement in camera trap data collection. Our results support the use of camera trap data where GPS collars and aerial surveys are not feasible and where higher-resolution data are needed for more localized management. Finally, we discuss our experience with two different types of camera traps to highlight the pros and cons of each approach.

Reference genome assembly and annotation of two Bacillus cereus sensu lato strains from Etosha National Park, Namibia.

Bacillus cereus sensu lato (s.l.) poses health and security issues. Here, we report the reference genome assembly of two Bacillus cereus s.l. strains, isolated from Etosha National Park, Namibia (FFI_BCgr36 and FFI_BCgr46). These unique genomes open for better understanding of environmental diversity and improvements in detection of threatening species.

Thinking with relations in nature conservation? A case study of the Etosha National Park and Haiǁom

AbstractThe area of the Etosha National Park in Namibia has been inhabited for many centuries by Haiǁom, a group of (now former) hunter‐gatherers. In 1907, Etosha was proclaimed as a game reserve, although Haiǁom were still allowed to live in the area until they were expelled in the 1950s due to then‐dominant ideas of fortress conservation. In recent years, Haiǁom have been provided with several resettlement farms by the Namibian government as a reaction to the colonial land dispossession. In this article, I explore the onto‐epistemology of Haiǁom (i.e. their being in and knowing the Etosha area), focusing on their relations with the land and with human and beyond‐the‐human beings before their eviction. I argue that the eviction implies not only economic marginalization but also social deprivation, which is inadequately addressed with resettlement. I suggest that thinking with relations, illustrated with the Haiǁom case, would call for other solutions in the context of measures taken for past land dispossessions and would open new paths for Namibia's nature conservation initiatives.

Variation in herbivore space use: comparing two savanna ecosystems with different anthrax outbreak patterns in southern Africa

BackgroundThe distribution of resources can affect animal range sizes, which in turn may alter infectious disease dynamics in heterogenous environments. The risk of pathogen exposure or the spatial extent of outbreaks may vary with host range size. This study examined the range sizes of herbivorous anthrax host species in two ecosystems and relationships between spatial movement behavior and patterns of disease outbreaks for a multi-host environmentally transmitted pathogen.MethodsWe examined range sizes for seven host species and the spatial extent of anthrax outbreaks in Etosha National Park, Namibia and Kruger National Park, South Africa, where the main host species and outbreak sizes differ. We evaluated host range sizes using the local convex hull method at different temporal scales, within-individual temporal range overlap, and relationships between ranging behavior and species contributions to anthrax cases in each park. We estimated the spatial extent of annual anthrax mortalities and evaluated whether the extent was correlated with case numbers of a given host species.ResultsRange size differences among species were not linearly related to anthrax case numbers. In Kruger the main host species had small range sizes and high range overlap, which may heighten exposure when outbreaks occur within their ranges. However, different patterns were observed in Etosha, where the main host species had large range sizes and relatively little overlap. The spatial extent of anthrax mortalities was similar between parks but less variable in Etosha than Kruger. In Kruger outbreaks varied from small local clusters to large areas and the spatial extent correlated with case numbers and species affected. Secondary host species contributed relatively few cases to outbreaks; however, for these species with large range sizes, case numbers positively correlated with outbreak extent.ConclusionsOur results provide new information on the spatiotemporal structuring of ranging movements of anthrax host species in two ecosystems. The results linking anthrax dynamics to host space use are correlative, yet suggest that, though partial and proximate, host range size and overlap may be contributing factors in outbreak characteristics for environmentally transmitted pathogens.

Fencing affects movement patterns of two large carnivores in Southern Africa

The erection of peripheral fences around protected areas has been shown to adversely affect the movement behavior, genetic connectivity and long-term viability of animal populations. However, fencing is also used extensively to reduce human encroachment, limit poaching activities, and mitigate human–wildlife conflicts. Limited information on the movement behavior of carnivores is available along the periphery of protected areas, particularly in semi-arid environments. To quantify the potential effects of the 824 km perimeter fence surrounding the Etosha National Park (Etosha, Namibia) on the movement and space use of carnivores, we collected GPS movement data from 36 lions (Panthera leo) and seven spotted hyenas (Crocuta crocuta) over a 14-year period. For these two apex carnivores, we measured the impact of the Etosha fence on their movements (step lengths and path straightness), calculated the likelihood of fence crossings, and evaluated the spatial–temporal characteristics of these crossings. Our results indicated that lions and hyenas moved faster (longer step lengths) when closer to the fence. Lions showed no variation in path straightness with respect to the fence but hyenas had straighter movement paths when closer to the fence. When moving within the vicinity of the fence, lions had a 9% likelihood of crossing the fence, while hyenas were much more likely (18%) to do so, which suggests that the Etosha fence was more permeable for hyenas than for lions. Fence crossings predominantly occurred at night for both species. Lions were more likely to cross the fence during the cold dry season (May–August) whereas hyenas crossed more often during the warm wet season (December–April). Female and adult lions had a higher probability of crossing the fence than males and subadults. These findings could be useful in the development of appropriate strategies to mitigate human–carnivore conflicts and thus promote the conservation of carnivores in multiple-use areas along the periphery of the park. Further research is needed to better elucidate the impact of perimeter fences on a broader suite of carnivores and other large mammals, including species-specific variation in the response to fences, as well as the economic benefits of fencing provided to local communities.

Development of a novel immobilisation protocol for black-faced impala (Aepyceros melampus ssp. petersi) in Etosha National Park.

Black-faced impala (Aepyceros melampus ssp. petersi) are endemic to Namibia where conservation management involves immobilisation and translocation, and mortality with current protocols is common. Critically evaluated field immobilisation protocols are needed to maximise animal safety. This prospective study was done in two phases: the first compared etorphine- and thiafentanil-based combinations, the second evaluated the influence of oxygen in impala receiving the thiafentanil-based combination. Animals (10 per group) received 50 mg ketamine (K) and 10 mg butorphanol (B), with either 2.0 mg etorphine (E) or 2.0 mg thiafentanil (T). A third group of ten impala were anaesthetised using TKB with supplemental nasal oxygen (O) at a rate of 5 L/minute. Behavioural, metabolic and physiological variables were assessed within five minutes of recumbency and at 10, 15, and 20 minutes post-recumbency. Statistical analyses for non-parametric data were performed to compare the treatment groups as well as time points; p ≤ 0.05 considered significant. Following darting, 7/10 EKB animals were standing when approached, compared to 2/20 in the thiafentanil treatment groups. Time to first effect was significantly higher for EKB (155 ± 105.7 seconds) compared to TKBO (61.5 ± 21.4 seconds). Time to sternal after darting was significantly higher with EKB (411.6 ± 174 seconds) compared to TKB (160.5 ± 85.4 seconds) and TKBO (166 ± 77.3 seconds). This study builds on previous work investigating the effects of potent opioids on impala and is the first evaluating their use in a field setting. The thiafentanil combination had a faster onset and resulted in a smoother induction than the etorphine combination. Additionally, oxygenation improved in animals receiving oxygen supplementation.

A Qualitative Exploration of Conflicts in Human-Wildlife Interactions in Namibia’s Kunene Region

Wildlife numbers are declining globally due to anthropogenic pressures. In Namibia, however, wildlife populations increased with policy instruments that allow private ownership and incentivize their sustainable use. Antithetically, this resulted in increased resource competition between humans and wildlife and triggered conflicts among various stakeholder groups. This paper summarizes the results of a qualitative exploration of conflicts in wildlife management in Namibia’s Kunene Region, adjacent to Etosha National Park. We conducted a workshop and expert interviews with stakeholders from relevant sectors. Our qualitative research sheds light on societal conflicts over wildlife that originate from diverging interests, livelihood strategies, moral values, knowledge holders, personal relations and views on institutional procedures. We frame our insights into conflicting human–wildlife interactions with theoretical concepts of social-ecological systems, ecosystem services and ecosystem disservices and open the floor for quantitative assessments. Overall, our results may present a suitable way of understanding biodiversity conflicts in a theoretical way.

The persistence of time: the lifespan of Bacillus anthracis spores in environmental reservoirs

Anthrax is a lethal bacterial zoonosis primarily affecting herbivorous wildlife and livestock. Upon host death Bacillus anthracis vegetative cells form spores capable of surviving for years in soil. Anthrax transmission requires host exposure to large spore doses. Thus, conditions that facilitate higher spore concentrations or promote spore survival will increase the probability that a pathogen reservoir infects future hosts. We investigated abiotic and pathogen genomic variation in relation to spore concentrations in surface soils (0–1 cm depth) at 40 plains zebra (Equus quagga) anthrax carcass sites in Namibia. Specifically, how initial spore concentrations and spore survival were affected by seasonality associated with the timing of host mortality, local soil characteristics, and pathogen genomic variation. Zebras dying of anthrax in wet seasons—the peak season for anthrax in Etosha National Park—had soil spore concentrations 1.36 orders of magnitude higher than those that died in dry seasons. No other variables considered affected spore concentrations, and spore survival rates did not differ among sites. Surface soils at these pathogen reservoirs remained culture positive for a range of 3.8–10.4 years after host death. Future research could evaluate if seasonal patterns in spore concentrations are driven by differences in sporulation success or levels of terminal bacteremia.

Flehmen, Osteophagia, and Other Behaviors of Giraffes (Giraffa giraffa angolensis): Vomeronasal Organ Adaptation.

The size of adult male giraffes (Giraffa giraffa angolensis) far exceeds the size of the females. At the Namutoni waterholes in Etosha National Park, bulls were seen many times each day screening adult females for their pending sexual receptivity by provoking them to urinate; this mainly involved sniffing their genitalia. If the female accedes to the male's invitation, she widens her hindleg stance, braces her body, and then urinates, usually for at least five seconds. The male places his muzzle and tongue in the urine stream, and then performs flehmen, often raising his head high in the air. Males never investigated urine on the ground. The bilateral papillae on the giraffe's hard palate connect with the nasopalatine ducts, which enter the bilateral vomeronasal organ (VNO). Unlike many mammals, the giraffe's VNO lacks a prominent connection to the nasal cavity and its connections to the oral cavity are primarily via the incisive papillae. Most days, some giraffes were observed searching for bones for extended periods, chewing them, and sometimes being troubled by a bone stuck in their mouth. A giraffe killed by lions was frequented for several days by a procession of giraffes investigating the carcass. A very dark-colored bull giraffe emitted loud pulsed growls that drove off most of the surrounding giraffes.

Africa’s drylands in a changing world: Challenges for wildlife conservation under climate and land-use changes in the Greater Etosha Landscape

Proclaimed in 1907, Etosha National Park in northern Namibia is an iconic dryland system with a rich history of wildlife conservation and research. A recent research symposium on wildlife conservation in the Greater Etosha Landscape (GEL) highlighted increased concern of how intensification of global change will affect wildlife conservation based on participant responses to a questionnaire. The GEL includes Etosha and surrounding areas, the latter divided by a veterinary fence into large, private farms to the south and communal areas of residential and farming land to the north. Here, we leverage our knowledge of this ecosystem to provide insight into the broader challenges facing wildlife conservation in this vulnerable dryland environment. We first look backward, summarizing the history of wildlife conservation and research trends in the GEL based on a literature review, providing a broad-scale understanding of the socioecological processes that drive dryland system dynamics. We then look forward, focusing on eight key areas of challenge and opportunity for this ecosystem: climate change, water availability and quality, vegetation and fire management, adaptability of wildlife populations, disease risk, human-wildlife conflict, wildlife crime, and human dimensions of wildlife conservation. Using this model system, we summarize key lessons and identify critical threats highlighting future research needs to support wildlife management. Research in the GEL has followed a trajectory seen elsewhere reflecting an increase in complexity and integration across biological scales over time. Yet, despite these trends, a gap exists between the scope of recent research efforts and the needs of wildlife conservation to adapt to climate and land-use changes. Given the complex nature of climate change, in addition to locally existing system stressors, a framework of forward-thinking adaptive management to address these challenges, supported by integrative and multidisciplinary research could be beneficial. One critical area for growth is to better integrate research and wildlife management across land-use types. Such efforts have the potential to support wildlife conservation efforts and human development goals, while building resilience against the impacts of climate change. While our conclusions reflect the specifics of the GEL ecosystem, they have direct relevance for other African dryland systems impacted by global change.

A framework for integrating inferred movement behavior into disease risk models

Movement behavior is an important contributor to habitat selection and its incorporation in disease risk models has been somewhat neglected. The habitat preferences of host individuals affect their probability of exposure to pathogens. If preference behavior can be incorporated in ecological niche models (ENMs) when data on pathogen distributions are available, then variation in such behavior may dramatically impact exposure risk. Here we use data from the anthrax endemic system of Etosha National Park, Namibia, to demonstrate how integrating inferred movement behavior alters the construction of disease risk maps. We used a Maximum Entropy (MaxEnt) model that associated soil, bioclimatic, and vegetation variables with the best available pathogen presence data collected at anthrax carcass sites to map areas of most likely Bacillus anthracis (the causative bacterium of anthrax) persistence. We then used a hidden Markov model (HMM) to distinguish foraging and non-foraging behavioral states along the movement tracks of nine zebra (Equus quagga) during the 2009 and 2010 anthrax seasons. The resulting tracks, decomposed on the basis of the inferred behavioral state, formed the basis of step-selection functions (SSFs) that used the MaxEnt output as a potential predictor variable. Our analyses revealed different risks of exposure during different zebra behavioral states, which were obscured when the full movement tracks were analyzed without consideration of the underlying behavioral states of individuals. Pathogen (or vector) distribution models may be misleading with regard to the actual risk faced by host animal populations when specific behavioral states are not explicitly accounted for in selection analyses. To more accurately evaluate exposure risk, especially in the case of environmentally transmitted pathogens, selection functions could be built for each identified behavioral state and then used to assess the comparative exposure risk across relevant states. The scale of data collection and analysis, however, introduces complexities and limitations for consideration when interpreting results.

Don't stop me now: Managed fence gaps could allow migratory ungulates to track dynamic resources and reduce fence related energy loss

In semi-arid environments characterized by erratic rainfall and scattered primary production, migratory movements are a key survival strategy of large herbivores to track resources over vast areas. Veterinary Cordon Fences (VCFs), intended to reduce wildlife-livestock disease transmission, fragment large parts of southern Africa and have limited the movements of large wild mammals for over 60 years. Consequently, wildlife-fence interactions are frequent and often result in perforations of the fence, mainly caused by elephants. Yet, we lack knowledge about at which times fences act as barriers, how fences directly alter the energy expenditure of native herbivores, and what the consequences of impermeability are. We studied 2-year ungulate movements in three common antelopes (springbok, kudu, eland) across a perforated part of Namibia's VCF separating a wildlife reserve and Etosha National Park using GPS telemetry, accelerometer measurements, and satellite imagery. We identified 2905 fence interaction events which we used to evaluate critical times of encounters and direct fence effects on energy expenditure. Using vegetation type-specific greenness dynamics, we quantified what animals gained in terms of high quality food resources from crossing the VCF. Our results show that the perforation of the VCF sustains herbivore-vegetation interactions in the savanna with its scattered resources. Fence permeability led to peaks in crossing numbers during the first flush of woody plants before the rain started. Kudu and eland often showed increased energy expenditure when crossing the fence. Energy expenditure was lowered during the frequent interactions of ungulates standing at the fence. We found no alteration of energy expenditure when springbok immediately found and crossed fence breaches. Our results indicate that constantly open gaps did not affect energy expenditure, while gaps with obstacles increased motion. Closing gaps may have confused ungulates and modified their intended movements. While browsing, sedentary kudu's use of space was less affected by the VCF; migratory, mixed-feeding springbok, and eland benefited from gaps by gaining forage quality and quantity after crossing. This highlights the importance of access to vast areas to allow ungulates to track vital vegetation patches.