High species diversity of Echinococcus spp. in wild mammals of Namibia.

An opportunistic survey for Echinococcus spp. in wild mammals was conducted in seven distinct study areas throughout Namibia, representing all major ecosystems, between 2012 and 2021. In total, 184 individually attributable faeces and 40 intestines were collected from eight species of carnivores, and 300 carcasses or organs of thirteen species of ungulates were examined for Echinococcus cysts. Nested PCR and sequencing of the mitochondrial nad1 gene led to the identification of five species of the Echinococcus granulosus sensu lato complex. Echinococcus canadensis G6/7 was found throughout Namibia at low frequency in lions, cheetahs, African wild dogs, black-backed jackals and oryx antelopes. Echinococcus equinus was present only in northern Namibia, locally at high frequency in lions, black-backed jackals and plains zebras. Echinococcus felidis was found only in one small area in the north-east of Namibia, but with high frequency in lions and warthogs. Echinococcus granulosus sensu stricto was identified only in two African wild dogs in the north-east of Namibia, and Echinococcus ortleppi occurred in central and southern Namibia in black-backed jackals and oryx antelopes. The development of fertile cysts indicated active intermediate host roles of oryx antelopes for E. canadensis and E. ortleppi, of warthogs for E. felidis, and of plains zebras for E. equinus. Our data support earlier hypotheses of exclusive or predominant wildlife life-cycles for E. felidis involving lions and warthogs, and - in Namibia - for E. equinus involving lions and/or black-backed jackals and plains zebras. Our data further support an interlink of wild and domestic transmission for E. ortleppi. A possible involvement of livestock and domestic dogs in transmission of E. canadensis G6/7 and E. granulosus s.s., the two parasite species with highest zoonotic potential, is uncertain for Namibia and needs further investigation.

GPS Telemetry Reveals a Zebra With Anthrax as Putative Cause of Death for Three Cheetahs in the Namib Desert.

Anthrax is a bacterial disease caused by Bacillus anthracis that affects wildlife, livestock and also humans in different parts of the world. It is endemic in some parts of Africa, including Namibia, with species differing in their susceptibility to the disease. Carnivores are typically less susceptible to anthrax than herbivores. Most carnivore species survive infection and have high seroprevalence against anthrax, whereas most herbivore species have low seroprevalence and typically die quickly when infected. Several reports have shown that cheetahs, unlike most other large carnivores, are susceptible to anthrax leading to a sudden death. This finding was suggested to be linked to the low genetic variability of cheetahs which might reduce an adequate immune response and thus explain such a high susceptibility to the disease. Here, we report an incidence of three free-ranging cheetahs that died within 24 h after feeding on a mountain zebra that tested positive for anthrax in the Namib Desert. We were able to reconstruct this incidence with the data recorded in the GPS (Global Positioning System) collar worn by one of the cheetahs and retrieved in the field. It is very likely that the cheetahs died from anthrax, although Bacillus anthracis could not be isolated from tissue and soil samples by bacterial culturing. The mountain zebra is the first described case of a wild animal that tested positive for anthrax in this arid area in southwestern of Namibia. We discuss the negative laboratory results of the cheetahs in the light of new insights of their immune system and its potential to mount a response against this bacteria.

Body size and digestive system shape resource selection by ungulates: A cross-taxa test of the forage maturation hypothesis.

The forage maturation hypothesis (FMH) states that energy intake for ungulates is maximised when forage biomass is at intermediate levels. Nevertheless, metabolic allometry and different digestive systems suggest that resource selection should vary across ungulate species. By combining GPS relocations with remotely sensed data on forage characteristics and surface water, we quantified the effect of body size and digestive system in determining movements of 30 populations of hindgut fermenters (equids) and ruminants across biomes. Selection for intermediate forage biomass was negatively related to body size, regardless of digestive system. Selection for proximity to surface water was stronger for equids relative to ruminants, regardless of body size. To be more generalisable, we suggest that the FMH explicitly incorporate contingencies in body size and digestive system, with small-bodied ruminants selecting more strongly for potential energy intake, and hindgut fermenters selecting more strongly for surface water.

Communication hubs of an asocial cat are the source of a human-carnivore conflict and key to its solution.

Human-wildlife conflicts occur worldwide. Although many nonlethal mitigation solutions are available, they rarely use the behavioral ecology of the conflict species to derive effective and long-lasting solutions. Here, we use a long-term study with 106 GPS-collared free-ranging cheetahs (Acinonyx jubatus) to demonstrate how new insights into the socio-spatial organization of this species provide the key for such a solution. GPS-collared territory holders marked and defended communication hubs (CHs) in the core area of their territories. The CHs/territories were distributed in a regular pattern across the landscape such that they were not contiguous with each other but separated by a surrounding matrix. They were kept in this way by successive territory holders, thus maintaining this overdispersed distribution. The CHs were also visited by nonterritorial cheetah males and females for information exchange, thus forming hotspots of cheetah activity and presence. We hypothesized that the CHs pose an increased predation risk to young calves for cattle farmers in Namibia. In an experimental approach, farmers shifted cattle herds away from the CHs during the calving season. This drastically reduced their calf losses by cheetahs because cheetahs did not follow the herds but instead preyed on naturally occurring local wildlife prey in the CHs. This implies that in the cheetah system, there are "problem areas," the CHs, rather than "problem individuals." The incorporation of the behavioral ecology of conflict species opens promising areas to search for solutions in other conflict species with nonhomogenous space use.

Sex and species differences of stress markers in sympatric cheetahs and leopards in Namibia.

Physiological stress markers may provide valuable insight for our understanding of costs of given life-history strategies or of wildlife health condition, most importantly in case of threatened species. In the last decade, there has been growing interest in the ecological relevance of cellular oxidative stress, which would provide complimentary information to that obtained by the classic analyses of glucocorticoid hormones. In this study, we analysed the sex and species variation of five blood-based markers of oxidative status, both molecular oxidative damage and antioxidant protection, in sympatric cheetahs (Acinonyx jubatus) and leopards (Panthera pardus) living on Namibian farmlands. Both these terrestrial carnivores are classified as vulnerable by the International Union for Conservation of Nature. We found that female cheetahs had significantly higher serum reactive oxygen metabolites of non-protein origin and lower glutathione peroxidase activity in whole blood than both male and female leopards and male cheetahs. We also found that cheetahs and leopards differed in the association between the two antioxidant enzymes glutathione peroxidase and superoxide dismutase. Correlations among oxidative status markers were stronger in female cheetahs than leopards or male cheetahs. Our results suggest that female cheetahs are more sensitive to local sources of stress. Our work did not corroborate the assumption that two species with different life histories consistently differ in key physiological traits.

Coping with intrasexual behavioral differences: Capture-recapture abundance estimation of male cheetah.

Population estimates are a fundamental requirement of ecology and conservation. While capture-recapture models are an established method for producing such estimates, their assumption of homogeneous capture probabilities is problematic given that heterogeneity in individual capture probability is inherent to most species. Such variation must be accounted for by abundance models; otherwise, biased estimates are risked.Here, we investigate the performance of four types of heterogeneity models for estimating abundance of male cheetah Acinonyx jubatus, a species with two distinct spatial tactics of territorial and nonterritorial (floater) males. The differences in spatial movements of territory holders and floaters are expected to result in intrasexual heterogeneous capture probabilities. Four heterogeneity models were used to model male abundance at five territories in central Namibia; (a) a spatial tactic model, (b) a finite mixture model, both run in program MARK, (c) a floater-only model, and (d) a heterogeneity Mh model, both run in the program CAPTURE. Camera trap data of cheetah, taken at frequently visited marking trees, were used to derive true abundance. Model results were compared to the true abundance to assess the accuracy of estimates.Only models (a), (b), and (c) were able to consistently produce accurate results. Mixture models do not require prior knowledge regarding spatial tactic of males, which might not always be available. Therefore, we recommend such models as the preferred model type for cheetahs.Results highlight the potential for mixture models in overcoming the challenges of capture probability heterogeneity and in particular their use with species where intrasexual behavioral differences exist.

Effects of host traits and land-use changes on the gut microbiota of the Namibian black-backed jackal (Canis mesomelas).

Host traits and environmental factors drive the natural variation in gut microbiota, and disruption in homeostasis can cause infections and chronic diseases. African wildlife is increasingly facing human-induced agricultural habitats, which also amplifies the contact probability with livestock with unknown consequences for wildlife gut microbiotas and the risk of transmission of potentially pathogenic bacteria. We applied high-throughput sequencing of bacterial 16S rRNA genes and microsatellite genotyping to investigate the impact of host traits and habitat use on the gut microbiotas of black-backed jackals (Canis mesomelas). This abundant carnivore inhabits livestock and game farms in central Namibia and is often persecuted as pathogen reservoir and vector. We further compared the gut microbiotas of black-backed jackals to other wild and domestic carnivores, herbivores and an omnivore, to disentangle the effects of environment, host species and dietary preference. In black-backed jackals, intrinsic host traits had a stronger impact in shaping the host-bacteria relationship than environmental factors. Nevertheless, the abundance of bacterial operational taxonomic units (OTUs) differed in individuals from livestock and game farms for specific bacterial genera such as Lactobacillus and Clostridium. We found, however, no evidence that black-backed jackals harbour abnormal levels of OTUs related to potential bacterial pathogens or that livestock farming has a negative impact on their health. We present here the first study investigating simultaneously the impact of host traits and environmental factors on gut microbiotas of a wildlife carnivore that occurs in a human-modified habitat.

The global decline of cheetah Acinonyx jubatus and what it means for conservation.

Establishing and maintaining protected areas (PAs) are key tools for biodiversity conservation. However, this approach is insufficient for many species, particularly those that are wide-ranging and sparse. The cheetah Acinonyx jubatus exemplifies such a species and faces extreme challenges to its survival. Here, we show that the global population is estimated at ∼7,100 individuals and confined to 9% of its historical distributional range. However, the majority of current range (77%) occurs outside of PAs, where the species faces multiple threats. Scenario modeling shows that, where growth rates are suppressed outside PAs, extinction rates increase rapidly as the proportion of population protected declines. Sensitivity analysis shows that growth rates within PAs have to be high if they are to compensate for declines outside. Susceptibility of cheetah to rapid decline is evidenced by recent rapid contraction in range, supporting an uplisting of the International Union for the Conservation of Nature (IUCN) Red List threat assessment to endangered. Our results are applicable to other protection-reliant species, which may be subject to systematic underestimation of threat when there is insufficient information outside PAs. Ultimately, conserving many of these species necessitates a paradigm shift in conservation toward a holistic approach that incentivizes protection and promotes sustainable human-wildlife coexistence across large multiple-use landscapes.

Immunogenetic variation and differential pathogen exposure in free-ranging cheetahs across Namibian farmlands.

BACKGROUND: Genes under selection provide ecologically important information useful for conservation issues. Major histocompatibility complex (MHC) class I and II genes are essential for the immune defence against pathogens from intracellular (e.g. viruses) and extracellular (e.g. helminths) origins, respectively. Serosurvey studies in Namibian cheetahs (Acinonyx juabuts) revealed higher exposure to viral pathogens in individuals from north-central than east-central regions. Here we examined whether the observed differences in exposure to viruses influence the patterns of genetic variation and differentiation at MHC loci in 88 free-ranging Namibian cheetahs. METHODOLOGY/PRINCIPAL FINDINGS: Genetic variation at MHC I and II loci was assessed through single-stranded conformation polymorphism (SSCP) analysis and sequencing. While the overall allelic diversity did not differ, we observed a high genetic differentiation at MHC class I loci between cheetahs from north-central and east-central Namibia. No such differentiation in MHC class II and neutral markers were found. CONCLUSIONS/SIGNIFICANCE: Our results suggest that MHC class I variation mirrors the variation in selection pressure imposed by viruses in free-ranging cheetahs across Namibian farmland. This is of high significance for future management and conservation programs of this species.

Diversity and evolutionary patterns of immune genes in free-ranging Namibian leopards (Panthera pardus pardus).

The genes of the major histocompatibility complex (MHC) are a key component of the mammalian immune system and have become important molecular markers for fitness-related genetic variation in wildlife populations. Currently, no information about the MHC sequence variation and constitution in African leopards exists. In this study, we isolated and characterized genetic variation at the adaptively most important region of MHC class I and MHC class II-DRB genes in 25 free-ranging African leopards from Namibia and investigated the mechanisms that generate and maintain MHC polymorphism in the species. Using single-stranded conformation polymorphism analysis and direct sequencing, we detected 6 MHC class I and 6 MHC class II-DRB sequences, which likely correspond to at least 3 MHC class I and 3 MHC class II-DRB loci. Amino acid sequence variation in both MHC classes was higher or similar in comparison to other reported felids. We found signatures of positive selection shaping the diversity of MHC class I and MHC class II-DRB loci during the evolutionary history of the species. A comparison of MHC class I and MHC class II-DRB sequences of the leopard to those of other felids revealed a trans-species mode of evolution. In addition, the evolutionary relationships of MHC class II-DRB sequences between African and Asian leopard subspecies are discussed.