In Africa, humans and large carnivores compete over access to resources, including prey. Disturbance by humans to kills made by carnivores, often for purposes of obtaining all or portions of the carcass, constitutes a form of human-wildlife conflict. However the occurrence of this practice, known as human kleptoparasitism, and its impact on carnivores has received little scientific attention. We obtained expert opinions from African lion researchers and stakeholders via a standardized questionnaire to characterize the geographic extent and frequency of human kleptoparasitism as it occurs in modern times. Our survey found modern human kleptoparasitism on kills made by lions, and possibly other large carnivores in Africa, to be geographically more widespread than previously reported. Meat lost to humans requires carnivores to hunt and kill additional prey thereby causing stress, increasing their energetic costs and risks of natural injury, and exposing them to risk of direct injury or death from human usurpers. Because of their conspicuous behaviors and tendency towards killing large-bodied prey, lions are particularly susceptible to humans detecting their kills. While human kleptoparasitism was geographically widespread, socio-economic factors influenced the frequency of occurrence. Prey type (wild game or domestic livestock) influenced human attitudes towards meat theft; ownership allows for legal recovery of livestock carcasses, while possessing wild game meat is mostly illegal and may incur penalties. Meat theft was associated with other illegal activities (i.e., illegal mining) and most prevalent among people of low income, including underpaid game scouts. Despite quantifiable costs to carnivores of human disturbance to their kills, the majority of experts surveyed reported a lack of knowledge on this practice. We propose that human disturbance at kills, especially loss of prey through human kleptoparasitism, constitutes an important anthropogenic threat that may seriously impact energy budgets of individual lions and other scavengers when meat and carcasses are removed from the ecosystem, and that the costs incurred by carnivores warrants further investigation.
Sub-Saharan Africa is under-represented in global biodiversity datasets, particularly regarding the impact of land use on species’ population abundances. Drawing on recent advances in expert elicitation to ensure data consistency, 200 experts were convened using a modified-Delphi process to estimate ‘intactness scores’: the remaining proportion of an ‘intact’ reference population of a species group in a particular land use, on a scale from 0 (no remaining individuals) to 1 (same abundance as the reference) and, in rare cases, to 2 (populations that thrive in human-modified landscapes). The resulting bii4africa dataset contains intactness scores representing terrestrial vertebrates (tetrapods: ±5,400 amphibians, reptiles, birds, mammals) and vascular plants (±45,000 forbs, graminoids, trees, shrubs) in sub-Saharan Africa across the region’s major land uses (urban, cropland, rangeland, plantation, protected, etc.) and intensities (e.g., large-scale vs smallholder cropland). This dataset was co-produced as part of the Biodiversity Intactness Index for Africa Project. Additional uses include assessing ecosystem condition; rectifying geographic/taxonomic biases in global biodiversity indicators and maps; and informing the Red List of Ecosystems.
Predictions obtained from wildlife population models may be prone to influences associated with the quality of the data available for parameterization. We assessed accuracy of furtaker-assigned sex of bobcats (Lynx rufus; n = 123) harvested during 2017-2018 across the western USA by comparing those data to genetically assigned sex, the latter of which we assumed was 100% accurate. We also compared the precision of individual age estimates obtained through cementum annuli analysis (CAA) of the canines and incisors of harvested bobcats (n = 151), where true age was unknown. Because cementum-line deposition may vary across populations and by sex, we hypothesized that environmental factors may affect precision of age estimates obtained via CAA. Sex of bobcats as reported by furtakers was accurate 82% of the time, and the direction of sex-assignment error was approximately equivalent. Canine teeth were more precise than incisors for estimating age, age estimates for male bobcats may be more precise than for females, and precision of age estimates may decrease with age. However, use of incisors corrected with known rates of error may be preferred in some instances, such as studies that require live capture and tooth extraction or where assignment to age class is sufficient. In addition, we found that sex, median age, and mean elevation may influence precision of age estimates, whereas mean precipitation and geographic location (latitude, longitude) were uninformative. Knowledge of different types of errors associated with sex identification and age estimation may be integrated into population modeling efforts based on management objectives.
The impact of snaring and human-wildlife conflict (HWC) on large carnivore populations is of growing concern, and yet few empirical data are available. Mortality is the metric most often used, but non-lethal injuries that impact fitness are also important threats. However, because non-lethal injuries to wild carnivores are difficult to detect, they have received little study. Using straightforward forensic examination of the skulls of trophy-hunted lions and leopards from Luangwa Valley (LV) and Greater Kafue Ecosystem (GKE), Zambia, we identified non-lethal injuries consisting of snare damage to teeth and shotgun pellets in skulls. Wire snare entanglement can cause permanent, diagnostic damage to carnivore teeth when individuals bite and pull on the wire. Shotguns are used by poachers, as well as during HWCs to drive off carnivores perceived as threats. Carnivores struck by shotgun pellets can suffer non-lethal, but potentially toxic injuries such as pellets embedded in their skulls. Because poaching and HWC are generally more prevalent near human settlements, we predicted a higher incidence of anthropogenic injuries to carnivores in Luangwa where the human population is larger and more concentrated along protected area edges than Kafue. Contrary to expectation, anthropogenic injuries were more prevalent among lions and leopards in Kafue than Luangwa. Notably, definitive evidence of snare entanglement greatly surpassed previous estimates for these regions. Overall, 37% (41 in 112) of adult male lions (29% in Luangwa, 45% in Kafue) and 22% (10 in 45) of adult male leopards (17% in Luangwa, 26% in Kafue) examined had survived being snared at some point in their lifetime. Among adult male lions, 27% (30 in 112) had old shotgun pellet injuries to their skulls. Our procedure of forensic examination of carnivore skulls and teeth, some of which can be applied to live-captured animals, allows for improved detection of cryptic, non-lethal anthropogenic injuries. Further, our methods represent a consistent and economical way to track changes in the frequency of such injuries over time and between regions, thereby providing a direct measure of the effectiveness of conservation programs that seek to reduce poaching and HWC.
Direct comparisons between historical and contemporary populations allow for detecting changes in genetic diversity through time and assessment of the impact of habitat fragmentation. Here, we determined the genetic architecture of both historical and modern lions to document changes in genetic diversity over the last century. We surveyed microsatellite and mitochondrial genome variation from 143 high-quality museum specimens of known provenance, allowing us to directly compare this information with data from several recently published nuclear and mitochondrial studies. Our results provide evidence for male-mediated gene flow and recent isolation of local subpopulations, likely due to habitat fragmentation. Nuclear markers showed a significant decrease in genetic diversity from the historical (H-E=0.833) to the modern (H-E=0.796) populations, whereas mitochondrial genetic diversity was maintained (H-d=0.98 for both). Although the historical population appears to have been panmictic based on nDNA data, hierarchical structure analysis identified four tiers of genetic structure in modern populations and was able to detect most sampling locations. Mitogenome analyses identified four clusters: Southern, Mixed, Eastern, and Western and were consistent between modern and historically sampled haplotypes. Within the last century, habitat fragmentation caused lion subpopulations to become more geographically isolated as human expansion changed the African landscape. This resulted in an increase in fine-scale nuclear genetic structure and loss of genetic diversity as lion subpopulations became more differentiated, whereas mitochondrial structure and diversity were maintained over time.
ABSTRACTReported effects of trophy harvest often are controversial. The subject is nuanced and many studies lack details necessary to place their results in context. Consequently, many studies are misunderstood or their conclusions misapplied. We propose that all dialogues about trophy hunting include a definition of how they use the term trophy, details of variables measured and why they were selected, and explanations of temporal and spatial scales employed. Only with these details can potential effects of trophy hunting be understood in context and used for management and policy decisions. © 2021 The Wildlife Society.
Determining the incidence and causes of craniodental damage in wild carnivores is often constrained by limited access to specimens with associated ecological data, such as prey type and abundance. We assessed dental condition and cranial injuries in lion, leopard, and spotted hyena in relation to prey and predator populations in Zambia's Luangwa Valley, where large prey are more abundant and lion and leopard more numerous, and the Greater Kafue Ecosystem, where smaller prey species are more prevalent and lion and leopard less common. In Luangwa, lions had significantly higher rates of tooth fracture, and blunt trauma injuries attributable to prey-handling, compared to Kafue lions. In contrast, leopards in both regions had similar rates of tooth wear and breakage. Overall, lions showed a significantly higher tooth fracture rate than leopards on a per tooth basis. Spotted hyenas had the highest rates of tooth wear and fracture among all three carnivores, and greatly exceeded previously recorded rates based on historical samples. Despite larger numbers of lion and leopard in Luangwa, there was no difference in incidence of intraspecific injuries between regions. These results are consistent with a greater abundance of large prey species, especially buffalo, in the diets of Luangwa lions, and previous work showing a reliance on smaller prey species in Kafue throughout the large carnivore guild.
ABSTRACT The Scramble for Africa in the late 1800s marked the beginning of increased human population growth in Africa. Here, we determined the genetic architecture of both historical and modern lions to identify changes in genetic diversity that occurred during this period of landscape and anthropogenic change. We surveyed microsatellite and mitochondrial genetic variation from 143 high-quality museum specimens of known provenance and combined them with data from recently published nuclear and mitochondrial studies. Analysis of variation at 9 microsatellites and 280 polymorphic mitogenome SNPs indicate the presence of male-mediated gene flow and recent isolation of local subpopulations, likely due to habitat fragmentation. Nuclear markers showed a significant decrease in genetic diversity from the historical (H E =0.833) to the modern (H E =0.796) populations, while mitochondrial genetic diversity was maintained (Hd=0.98 for both). While the historical population appears to have been panmictic based on nDNA data, hierarchical structure analysis identified four tiers of fine structure in modern populations, able to detect most sampling locations. Mitochondrial analyses identified 4 clusters: Southern, Mixed, Eastern, and Western; and were consistent between modern and historically sampled haplotypes. Within the last century, habitat fragmentation caused lion subpopulations to become more isolated as human expansion changed the African landscape. This resulted in an increase in fine-scale nuclear genetic structure and loss of genetic diversity as subpopulations became more differentiated, while mitochondrial structure and diversity was maintained over time.
The Luangwa Valley in eastern Zambia is a transverse offshoot of the Great Rift Valley system. This region appears to have an isolating effect as evidenced by suspected endemic subspecies, such as the Cookson's wildebeest and Thornicroft's giraffe. Recent mitochondrial DNA studies demonstrated that African lions in Zambia consist of two highly diverse eastern and western sub-populations. Herein, we report nuclear and mitochondrial DNA results from 409 lions that support this population substructure across Zambia but proposes only partial isolation of the Luangwa Valley with more movement between the populations than previously thought. Population assignment analysis identifies two populations with little evidence of admixture assigning lions to either the eastern or western sub-populations. A high occurrence of private alleles and clear evidence for a Wahlund effect further justify the presence of a highly structured population. But, while mitochondrial DNA analysis still shows little to no matrilineal gene flow (F-ST = 0.53) between sub-populations, microsatellite analysis suggests there is gene flow (F-ST = 0.04) with low but significant isolation-by-distance and an average of 6 migrants per generation. Evidence of isolation-by-distance is also found in factorial correspondence analysis with the Lower Zambezi National Park and eastern corridor clusters overlapping isolated clusters of the Luangwa Valley and western subpopulation. From this evidence, the Luangwa Valley appears separated from the western sub-population with some dispersal through the southern regions of the eastern sub-population. Both the eastern and western sub-populations have high heterozygosity (0.68 and 0.69, respectively) and genetic diversity (0.47 and 0.50, respectively) values, indicative of genetically healthy populations.
A summary report and photographic catalogue of individually identified African wild dogs Lycaon pictus and their pack affiliations was compiled from sightings made during the period 2007-2012 in the southern Kafue ecosystem, Zambia. Ten sightings with photographs and/or videos were made first-hand by the authors while 12 additional sightings accompanied by photographs and/or videos were obtained from other sources. An additional seven verified sighting reports (lacking photographs) were included in a map of sighting distribution. A minimum of 90 African wild dogs in eight packs were documented. Mean pack size was X±SE=8.7±1.2 adult dogs/pack (range 4-13 adults, n=7 packs). Five litters were documented; one additional litter was inferred through observations of a heavily pregnant female. This study addresses a data gap on wild dogs in the southern Kafue and may assist conservation efforts for African wild dogs now underway in the greater Kafue ecosystem and larger Kavango Zambezi Transfrontier Conservation Area.
The biodiversity working group of the Arctic Council has developed pan-Arctic biodiversity monitoring plans to improve our ability to detect, understand and report on long-term change in Arctic biodiversity. The Arctic fox (Vulpes lagopus) was identified as a target of future monitoring because of its circumpolar distribution, ecological importance and reliance on Arctic ecosystems. We provide the first exhaustive survey of contemporary Arctic fox monitoring programmes, describing 34 projects located in eight countries. Monitored populations covered equally the four climate zones of the species’ distribution, and there were large differences between populations in long-term trends, multi-annual fluctuations, diet composition, degree of competition with red fox and human interferences. Den density, number of active dens, number of breeding dens and litter size were assessed in almost all populations, while projects varied greatly with respect to monitoring of other variables indicative of population status, ecosystem state or ecosystem function. We review the benefits, opportunities and challenges to increased integration of monitoring projects. We argue that better harmonizing protocols of data collection and data management would allow new questions to be addressed while adding tremendous value to individual projects. However, despite many opportunities, challenges remain. We offer six recommendations that represent decisive progress toward a better integration of Arctic fox monitoring projects. Further, our work serves as a template that can be used to integrate monitoring efforts of other species, thereby providing a key step for future assessments of global biodiversity.
Improved age estimation of African lions Panthera leo is needed to address a number of pressing conservation issues. Here we present a formula for estimating lion age to within six months of known age based on measuring the extent of pulp closure from X-rays, or Ratio Of tooth AReas (ROAR). Derived from measurements taken from lions aged 3-13 years for which exact ages were known, the formula explains 92% of the total variance. The method of calculating the pulp/ tooth area ratio, which has been used extensively in forensic science, is novel in the study of lion aging. As a quantifiable measure, ROAR offers improved lion age estimates for population modeling and investigations of age-related mortality, and may assist national and international wildlife authorities in judging compliance with regulatory measures involving age.
The Arctic fox (Vulpes lagopus) is a small canid with a circumpolar Arctic distribution. Several subspecies are recognized, including a subspecies known as the Pribilof fox (V. I. pribilofensis) endemic to the Pribilof Islands of Alaska, USA. Pribilof fox tissues were collected from the islands of St. Paul (n = 38) and St. George (n = 13). Levels of persistent organic pollutants (POPs) and stable isotopes of carbon and nitrogen were measured and the findings related to sex, age class, island and access to anthropogenic food resources using ANOVA and principal component analysis. The rank order for POPs in fat was polychlorinated biphenyls (Sigma PCBs) > chlordanes (Sigma CHLs) >> hexachlorocyclohexanes (Sigma HCHs) > DDTs (Sigma DDTs) > hexachlorobenzene (HCB) similar to polybrominated diphenyl ethers (SPBDEs). Adult females had lower mean levels of most POPs (Sigma PCBs, Sigma CHLs, Sigma HCHs, Sigma DDTs) and lower delta N-15 values than adult males. Foxes on St. Paul had significantly higher levels of most POPs than those on St. George, though St. George foxes were significantly higher in HCB. Foxes with high probability of access to anthropogenic foods had significantly lower levels of Sigma DDTs and lower delta N-15 values than foxes with a low probability of access. The observed differences in contaminant and stable isotope levels were consistent with fox use patterns of different food resources. POP concentrations in the tissues of some Pribilof foxes, especially from St. Paul, were higher than those associated with thresholds for adverse health effects. POPs may therefore be a factor for consideration in the conservation of Pribilof foxes.
Trophy hunting plays a significant role in wildlife conservation in some contexts in various parts of the world. Yet excessive hunting is contributing to species declines, especially for large carnivores. Simulation models suggest that sustainable hunting of African lions may be achieved by restricting offtakes to males old enough to have reared a cohort of offspring. We tested and expanded criteria for an age-based approach for sustainably regulating lion hunting. Using photos of 228 known-age males from ten sites across Africa, we measured change in ten phenotypic traits with age and found four age classes with distinct characteristics: 1–2.9years, 3–4.9years, 5–6.9years, and ≥7years. We tested the aging accuracy of professional hunters and inexperienced observers before and after training on aging. Before training, hunters accurately aged more lion photos (63%) than inexperienced observers (48%); after training, both groups improved (67–69%). Hunters overestimated 22% of lions <5years as 5–6.9years (unsustainable) but only 4% of lions <5years as ≥7years (sustainable). Due to the lower aging error for males ≥7years, we recommend 7years as a practical minimum age for hunting male lions. Results indicate that age-based hunting is feasible for sustainably managing threatened and economically significant species such as the lion, but must be guided by rigorous training, strict monitoring of compliance and error, and conservative quotas. Our study furthermore demonstrates methods for identifying traits to age individuals, information that is critical for estimating demographic parameters underlying management and conservation of age-structured species.
Countries that allow sport hunting of African lions, Panthera leo , increasingly are mandating minimum age regulations for harvested individuals that require accurate aging techniques. However, individual variation in dental characteristics used for aging could result in errors and has not been quantified. We assessed individual variation in three dental methods for estimating age of lions using left and right paired PM 2 s from 31 free‐ranging lions of unknown age from Zambia. Assignment of individuals to age class based on crown wear was largely consistent (87%) between paired PM 2 s; measures of pulp chamber closure also were highly consistent (< 4% variation) between pairs. Though pulp chamber closure varied little between pairs, calibrating pulp closure with known‐aged lions is needed to fully develop this method. In contrast, cementum line counts differed between paired PM 2 s in 94% of lions, ranging from 1 to 7 cementum lines (i.e. 0.5–3.5 years). Of lions with cementum lion counts that differed between teeth, 45% were estimated as meeting the 5‐ or 6‐year‐old minimum age used in trophy lion monitoring programs with one PM 2 , but not the paired PM 2 . Though individual variation in cementum line counts may render this technique unsuitable for aging lions, using the maximum or average count from both PM 2 s warrants further investigation. Individual variation in dental characteristics should be considered when developing lion aging techniques based on dentition.
Sport hunting has reportedly multiple benefits to economies and local communities; however, few of these benefits have been quantified. As part of their lease agreements with the Zambia Wildlife Authority, sport hunting operators in Zambia are required to provide annually to local communities free of charge i.e., provision a percentage of the meat obtained through sport hunting. We characterized provisioning of game meat to rural communities by the sport hunting industry in Zambia for three game management areas (GMAs) during 2004-2011. Rural communities located within GMAs where sport hunting occurred received on average > 6,000 kgs per GMA of fresh game meat annually from hunting operators. To assess hunting industry compliance, we also compared the amount of meat expected as per the lease agreements versus observed amounts of meat provisioned from three GMAs during 2007-2009. In seven of eight annual comparisons of these GMAs, provisioning of meat exceeded what was required in the lease agreements. Provisioning occurred throughout the hunting season and peaked during the end of the dry season (September-October) coincident with when rural Zambians are most likely to encounter food shortages. We extrapolated our results across all GMAs and estimated 129,771 kgs of fresh game meat provisioned annually by the sport hunting industry to rural communities in Zambia at an approximate value for the meat alone of > US$600,000 exclusive of distribution costs. During the hunting moratorium (2013-2014), this supply of meat has halted, likely adversely affecting rural communities previously reliant on this food source. Proposed alternatives to sport hunting should consider protein provisioning in addition to other benefits (e.g., employment, community pledges, anti-poaching funds) that rural Zambian communities receive from the sport hunting industry.
Analysis of DNA sequence diversity at the 12S to 16S mitochondrial genes of 165 African lions (Panthera leo) from five main areas in Zambia has uncovered haplotypes which link Southern Africa with East Africa. Phylogenetic analysis suggests Zambia may serve as a bridge connecting the lion populations in southern Africa to eastern Africa, supporting earlier hypotheses that eastern-southern Africa may represent the evolutionary cradle for the species. Overall gene diversity throughout the Zambian lion population was 0.7319 +/-0.0174 with eight haplotypes found; three haplotypes previously described and the remaining five novel. The addition of these five novel haplotypes, so far only found within Zambia, nearly doubles the number of haplotypes previously reported for any given geographic location of wild lions. However, based on an AMOVA analysis of these haplotypes, there is little to no matrilineal gene flow (FST = 0.47) when the eastern and western regions of Zambia are considered as two regional sub-populations. Crossover haplotypes (H9, H11, and Z1) appear in both populations as rare in one but common in the other. This pattern is a possible result of the lion mating system in which predominately males disperse, as all individuals with crossover haplotypes were male. The determination and characterization of lion sub-populations, such as done in this study for Zambia, represent a higher-resolution of knowledge regarding both the genetic health and connectivity of lion populations, which can serve to inform conservation and management of this iconic species.
The evolutionary history of a species is key for understanding the taxonomy and for the design of effective management strategies for species conservation. The knowledge about the phylogenetic position of the lion (Panthera leo) in West/Central Africa is largely based on mitochondrial markers. Previous studies using mtDNA only have shown this region to hold a distinct evolutionary lineage. In addition, anthropogenic factors have led to a strong decline in West/Central African lion numbers, thus, the conservation value of these populations is particularly high. Here, we investigate whether autosomal markers are concordant with previously described phylogeographic patterns, and confirm the unique position of the West/Central African lion. Analysis of 20 microsatellites and 1,454 bp of the mitochondrial DNA in 16 lion populations representing the entire geographic range of the species found congruence in both types of markers, identifying four clusters: 1) West/Central Africa, 2) East Africa, 3) Southern Africa and 4) India. This is not in line with the current taxonomy, as defined by the IUCN, which only recognizes an African and an Asiatic subspecies. There are no indications that genetic diversity in West/Central Africa lions is lower than in either East or Southern Africa, however, given this genetic distinction and the recent declines of lion numbers in this region, we strongly recommend prioritization of conservation projects in West/Central Africa. As the current taxonomic nomenclature does not reflect the evolutionary history of the lion, we suggest that a taxonomic revision of the lion is warranted.