Creating Larger and Better Connected Protected Areas Enhances the Persistence of Big Game Species in the Maputaland-Pondoland-Albany Biodiversity Hotspot

The ideal conservation planning approach would enable decision-makers to use population viability analysis to assess the effects of management strategies and threats on all species at the landscape level. However, the lack of high-quality data derived from long-term studies, and uncertainty in model parameters and/or structure, often limit the use of population models to only a few species of conservation concern. We used spatially explicit metapopulation models in conjunction with multi-criteria decision analysis to assess how species-specific threats and management interventions would affect the persistence of African wild dog, black rhino, cheetah, elephant, leopard and lion, under six reserve scenarios, thereby providing the basis for deciding on a best course of conservation action in the South African province of KwaZulu-Natal, which forms the central component of the Maputaland-Pondoland-Albany biodiversity hotspot. Overall, the results suggest that current strategies of managing populations within individual, small, fenced reserves are unlikely to enhance metapopulation persistence should catastrophic events affect populations in the future. Creating larger and better-connected protected areas would ensure that threats can be better mitigated in the future for both African wild dog and leopard, which can disperse naturally, and black rhino, cheetah, elephant, and lion, which are constrained by electric fences but can be managed using translocation. The importance of both size and connectivity should inform endangered megafauna conservation and management, especially in the context of restoration efforts in increasingly human-dominated landscapes.     

Conservation of severely fragmented populations: lessons from the transformation of uncoordinated reintroductions of cheetahs (<Emphasis Type="Italic">Acinonyx jubatus</Emphasis>) into a managed metapopulation with self-sustained growth

We document and evaluate the use of metapopulation management to conserve a declining population of 217 cheetahs in 40 subpopulations. Metapopulation management resembles a natural metapopulation, but dispersal success, demographic rescue effects and genetic viability are enhanced by moving suitable individuals to selected habitat fragments. Unfortunately, history and results of metapopulation management are rarely published. Cheetahs, extirpated from 85% of South Africa, were reintroduced from Namibian and South African ranches into fenced reserves. During 1965–2009 343 cheetahs were reintroduced, yet reserves held only 289 in 2009. Then translocations of free-roaming cheetahs were halted, and numbers dropped to 217 on 40 reserves by 2012. A metapopulation project was launched, and key conservation problems indentified from interviews and records. Thirty-five percent of reserves had no breeding cheetahs, 13% were inbreeding, fence quality was erratic, 3% of cheetahs were sold into captivity annually, and 28% of cheetah mortalities were anthropogenic. Lions accounted for 31% of mortality, perhaps elevated by lion-inexperienced cheetahs and high lion densities. These problems were addressed, and cheetahs were translocated between reserves. Although the median reserve size was only 125 km² holding four cheetahs, and 80% of reserves were privately owned, in 6 years the metapopulation grew by 51% to 328 cheetahs on 51 reserves, while genetic diversity was managed and monitored. Thus, using metapopulation management, low density species and associated key processes, including carnivory or mega-herbivory, can be conserved in relatively small reserves in regions with dense human populations precluding  natural gene flow.     

Conserving large carnivores: dollars and fence

Conservationists often advocate for landscape approaches to wildlife management while others argue for physical separation between protected species and human communities, but direct empirical comparisons of these alternatives are scarce. We relate African lion population densities and population trends to contrasting management practices across 42 sites in 11 countries. Lion populations in fenced reserves are significantly closer to their estimated carrying capacities than unfenced populations. Whereas fenced reserves can maintain lions at 80% of their potential densities on annual management budgets of $500 km⁻², unfenced populations require budgets in excess of $2000 km⁻² to attain half their potential densities. Lions in fenced reserves are primarily limited by density dependence, but lions in unfenced reserves are highly sensitive to human population densities in surrounding communities, and unfenced populations are frequently subjected to density‐independent factors. Nearly half the unfenced lion populations may decline to near extinction over the next 20–40 years.     

MtDNA haplotypes, sequence divergence, and morphological variation in Gray-breasted Wood Wrens (Henichorina leucophrys) and their conservation implications

ABSTRACT.   To reveal more clearly the places that are critical for generating and sustaining species diversity, the distribution of genetic diversity is informative for conservation. Evolutionary significant units (ESUs) represent distinct lineages within a species and are defined by genetic and morphological differences. We examine morphometrics and mitochondrial DNA sequences of two coastal and three Andean populations of Gray-breasted Wood Wrens ( Henicorhina leucophrys ) in Ecuador to determine if any populations are ESUs. Coastal Wood Wrens diverged genetically by 2.4%, 3.5%, and 4.4% from Andean populations of the northwest (Bellavista), southwest (Sural-Caucha), and southeast (Romerillos), respectively. The five populations (34 individuals) exhibited nine haplotypes. Both genetic and ecological results indicate that coastal Wood Wrens found in the Colonche Hills are ESUs, as are the Andean populations. In contrast with temperate regions, where large reserves connected by corridors may be effective, preservation of biodiversity in tropical South America may require protection of more localized areas.     

Identification of evolutionarily significant units in the Cuban endemic damselfly <Emphasis Type="Italic">Hypolestes trinitatis</Emphasis> (Odonata: Hypolestidae)

Species classification may not reflect the underlying/cryptic genetic diversity which should otherwise be conserved as it represents the potential of populations to evolve and adapt. The identification of evolutionarily significant units (ESUs) allows cryptic genetic diversity to be taken into account when designating conservation priorities. Here, we used mitochondrial and nuclear DNA sequences integrated with ecological niche models (ENM) to identify ESUs in Hypolestes trinitatis, a threatened Cuban endemic damselfly species. We found that this species comprises two distinct genetic groups in Central and Eastern Cuba respectively, which are also geographically isolated, as shown by ENM. Therefore, we propose these groups to be considered as different ESUs. According with their extent of occurrence, number of locations and inferred decline of habitat extent and quality, Central and Eastern ESUs qualify as Endangered [EN B1b(iii)] and Vulnerable [VU B1b(iii)], respectively.     

Genetic diversity,evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa

Aim In recent decades there has been a marked decline in the numbers of African lions (Panthera leo), especially in West Africa where the species is regionally endangered. Based on the climatological history of western Africa, we hypothesize that West and Central African lions have a unique evolutionary history, which is reflected by their genetic makeup. Location Sub‐Saharan Africa and India, with special focus on West and Central Africa. Method In this study 126 samples, throughout the lion’s complete geographic range, were subjected to phylogenetic analyses. DNA sequences of a mitochondrial region, containing cytochrome b, tRNAPro, tRNAThr and the left part of the control region, were analysed. Results Bayesian, maximum likelihood and maximum parsimony analyses consistently showed a distinction between lions from West and Central Africa and lions from southern and East Africa. West and Central African lions are more closely related to Asiatic lions than to the southern and East African lions. This can be explained by a Pleistocene extinction and subsequent recolonization of West Africa from refugia in the Middle East. This is further supported by the fact that the West and Central African clade shows relatively little genetic diversity and is therefore thought to be an evolutionarily young clade. Main conclusions The taxonomic division between an African and an Asian subspecies does not fully reflect the overall genetic diversity within lions. In order to conserve genetic diversity within the species, genetically distinct lineages should be prioritized. Understanding the geographic pattern of genetic diversity is key to developing conservation strategies, both for in situ management and for breeding of captive stocks.     

Extraordinary micro-endemism in Australian desert spring amphipods

Increasing pressure for water in the Australian arid zone is placing enormous stress on the diverse endemic communities inhabiting desert springs. Detailed information about the evolutionary processes occurring within and between individual endemic species will help to develop effective and biologically relevant management strategies this fragile ecosystem. To help determine conservation priorities, we documented the genetic structure of the endemic freshwater amphipod populations in springs fed by the Great Artesian Basin in central Australia. Phylogenetic and phylogeographic history and genetic diversity measures were examined using nuclear and mitochondrial DNA from approximately 500 chiltoniid amphipods across an entire group of springs. Pronounced genetic diversity was identified, demonstrating that levels of endemism have been grossly underestimated in these amphipods. Using the GMYC model, 13 genetically divergent lineages were recognized as Evolutionarily Significant Units (ESUs), all of which could be considered as separate species. The results show that due to the highly fragmented ecosystem, these taxa have highly restricted distributions. Many of the identified ESUs are endemic to a very small number of already degraded springs, with the rarest existing in single springs. Despite their extraordinarily small ranges, most ESUs showed relative demographic stability and high levels of genetic diversity, and genetic diversity was not directly linked to habitat extent. The relatively robust genetic health of ESUs does not preclude them from endangerment, as their limited distributions ensure they will be highly vulnerable to future water extraction.     

The Lion in West Africa Is Critically Endangered

The African lion has declined to <35,000 individuals occupying 25% of its historic range. The situation is most critical for the geographically isolated populations in West Africa, where the species is considered regionally endangered. Elevating their conservation significance, recent molecular studies establish the genetic distinctiveness of West and Central African lions from other extant African populations. Interventions to save West African lions are urgently required. However formulating effective conservation strategies has been hampered by a lack of data on the species'' current distribution, status, and potential management deficiencies of protected areas (PAs) harboring lions. Our study synthesized available expert opinion and field data to close this knowledge gap, and formulate recommendations for the conservation of West African lions. We undertook lion surveys in 13 large (>500 km²) PAs and compiled evidence of lion presence/absence for a further eight PAs. All PAs were situated within Lion Conservation Units, geographical units designated as priority lion areas by wildlife experts at a regional lion conservation workshop in 2005. Lions were confirmed in only 4 PAs, and our results suggest that only 406 (273–605) lions remain in West Africa, representing <250 mature individuals. Confirmed lion range is estimated at 49,000 km², or 1.1% of historical range in West Africa. PAs retaining lions were larger than PAs without lions and had significantly higher management budgets. We encourage revision of lion taxonomy, to recognize the genetic distinctiveness of West African lions and highlight their potentially unique conservation value. Further, we call for listing of the lion as critically endangered in West Africa, under criterion C2a(ii) for populations with <250 mature individuals. Finally, considering the relative poverty of lion range states in West Africa, we call for urgent mobilization of investment from the international community to assist range states to increase management effectiveness of PAs retaining lions.     

Differences in woody vegetation are unrelated to use by African elephants (Loxodonta africana) in Mkhuze Game Reserve, South Africa

The reintroduction of African elephants into fenced game reserves throughout South Africa has presented managers with several challenges. Although elephants are a natural part of southern African ecosystems, their confinement to fenced protected areas in South Africa has exacerbated their potential to impact their habitats negatively. However, many studies investigating the impact of elephants have failed to control for the effects of other browsers on the vegetative community. In this study, we used location data on an elephant herd to delineate high-use and low-use areas. This paired design allowed us to minimize confounding factors that could explain differences in the structure, diversity and utilization of woody species. We found little evidence to suggest elephant-mediated change in, or selection for, the structure or diversity of woody species; however, our results suggest that elephants may be altering the composition of species by preferentially using areas with higher canopy diversity and by enhancing sapling recruitment. Although stripping of bark was higher in high-use areas, there was no evidence of differential mortality of tree species. Therefore, in our study area, and over the current time scale, elephants are having a negligible impact on the vegetative community.     

Phylogeographic evidence links the threatened ‘Grampians’ Mountain Dragon (Rankinia diemensis Grampians) with Tasmanian populations: conservation implications in south-eastern Australia

The importance of protecting genetic diversity within a species is increasingly being recognised by conservation management authorities. However, discrepancies in conservation policy between authorities, such as state versus national bodies, can have significant implications for species management when they cross state boundaries. We conducted a phylogeographic study of the south-eastern Australian lizard Rankinia diemensis to identify evolutionary significant units (ESUs), including the endangered population from the Grampians National Park in western Victoria. Phylogenetic analyses of two gene regions (mtDNA: ND2; nuclear: RAG1) revealed high levels of genetic divergence between populations, indicating isolation over long evolutionary time frames. Based on criteria of genetic divergence and isolation, R. diemensis contains at least two ESUs that require specific management. We found that R. diemensis from the Grampians are closely related to Tasmanian populations, but that the divergence between these regions is great enough (3.7 % mtDNA) that they should be considered separate ESUs. However, we believe the close evolutionary ties between these two regions needs to be taken into account; yet under current practises, conservation management of subspecific ESUs relies on state-level efforts. We argue that another population that occurs on the Victorian coast also qualifies as an ESU and requires targeted conservation action. Rankinia diemensis provides a case-in-point of the discrepancy between the state-level approach of maintaining genetic variation within a species and the more conservative Commonwealth focus on conserving biodiversity at the species level.     

The origin, current diversity and future conservation of the modern lion (Panthera leo)

Understanding the phylogeographic processes affecting endangered species is crucial both to interpreting their evolutionary history and to the establishment of conservation strategies. Lions provide a key opportunity to explore such processes; however, a lack of genetic diversity and shortage of suitable samples has until now hindered such investigation. We used mitochondrial control region DNA (mtDNA) sequences to investigate the phylogeographic history of modern lions, using samples from across their entire range. We find the sub-Saharan African lions are basal among modern lions, supporting a single African origin model of modern lion evolution, equivalent to the 'recent African origin' model of modern human evolution. We also find the greatest variety of mtDNA haplotypes in the centre of Africa, which may be due to the distribution of physical barriers and continental-scale habitat changes caused by Pleistocene glacial oscillations. Our results suggest that the modern lion may currently consist of three geographic populations on the basis of their recent evolutionary history: North African-Asian, southern African and middle African. Future conservation strategies should take these evolutionary subdivisions into consideration.     

Identifying evolutionarily significant units for conservation of the endangered Malus sieversii using genome‐wide RADseq data

Malus sieversii, a wild progenitor of the domesticated apple, is an endangered species and is assigned second conservation priority by the China Plant Red Data Book. It is urgent to carry out in situ conservation of this species, but previous studies have not identified evolutionarily significant units (ESUs) for conservation management. In this study, we investigated the genetic diversity and relationships of six M. sieversii populations from China using integrated analysis of microsatellite (nSSR) data, genome‐wide SNPs and previous results in order to propose a reasonable conservation management. The results showed that levels of genetic diversity were inconsistently reflected by our nSSR and previous studies, suggesting that indices of genetic diversity are not effective to identify priority conservation areas for M. sieversii. Based on the selection criteria of ESUs for endangered species conservation, ESUs should reflect lineage divergence, geographical separation and different adaptive variation. Our phylogenetic tree based on genome‐wide SNPs yielded a clear relationship of divergent lineages among M. sieversii populations, leading to new different from those of previous studies. Three independent lineages, including the pairs of populations Huocheng‐Yining, Gongliu‐Xinyuan and Tuoli‐Emin, were identified. The geographic distances between populations among the different phylogenetic lineages were much greater than those within the same phylogenetic lineage. A cluster analysis on environmental variables showed that the three independent lineages inhabit different environmental conditions, suggesting that they may have adapted to different environments. Based on the results, we propose that three independent ESUs should be recognized as conservation units for M. sieversii in China.     

Evolutionarily significant units of the critically endangered leaf frog Pithecopus ayeaye (Anura,Phyllomedusidae) are not effectively preserved by the Brazilian protected areas network

Protected areas (PAs) are essential for biodiversity conservation, but their coverage is considered inefficient for the preservation of all species. Many species are subdivided into evolutionarily significant units (ESUs) and the effectiveness of PAs in protecting them needs to be investigated. We evaluated the usefulness of the Brazilian PAs network in protecting ESUs of the critically endangered Pithecopus ayeaye through ongoing climate change. This species occurs in a threatened mountaintop ecosystem known as campos rupestres. We used multilocus DNA sequences to delimit geographic clusters, which were further validated as ESUs with a coalescent approach. Ecological niche modeling was used to estimate spatial changes in ESUs’ potential distributions, and a gap analysis was carried out to evaluate the effectiveness of the Brazilian PAs network to protect P. ayeaye in the face of climate changes. We tested the niche overlap between ESUs to gain insights for potential management alternatives for the species. Pithecopus ayeaye contains at least three ESUs isolated in distinct mountain regions, and one of them is not protected by any PA. There are no climatic niche differences between the units, and only 4% of the suitable potential area of the species is protected in present and future projections. The current PAs are not effective in preserving the intraspecific diversity of P. ayeaye in its present and future range distributions. The genetic structure of P. ayeaye could represent a typical pattern in campos rupestres endemics, which should be considered for evaluating its conservation status.     

Fence management and time since pack formation influence African wild dog escapes from protected areas in South Africa

In human-dominated and highly fragmented landscapes, keeping wildlife within reserve boundaries is vital for conservation success. In South Africa, fences are a widely employed conservation management tool for protected areas and are successful in mitigating human-wildlife conflict. However, fences are permeable, and predators are able to cross through reserve fences. African wild dogs (Lycaon pictus) often leave fenced boundaries, resulting in high capture and translocation costs. Moreover, when wild dog packs (up to 30 individuals) leave fenced reserves they enter human-dominated landscapes where they face strong persecution and livestock predation incurs high costs. The factors driving packs to leave managed reserves are poorly understood, thus, to effectively manage wild dogs in fenced systems, it is important to understand why they leave reserve boundaries. There are several hypotheses as to why wild dogs cross through reserve fences, including inter- and intra-specific competition, social behaviour, management, prey density and environmental variability. Using a long-term dataset comprising 32 resident packs across five reserves, we investigated the relative strength of these hypotheses on the probability of wild dogs exiting a fenced reserve. During the 14-year study period, we recorded 154 exit events. We found that the interaction of fence integrity and time since pack formation were the primary factors affecting the probability of a pack leaving a reserve. When fence integrity was poor, escape probability decreased with pack age likely due to the exploratory behaviour of new packs. When fence integrity was average, escape probability increased with pack age likely due to the fitness benefits of holding larger and more exclusive territories as packs age. When fence integrity was good, the probability of a pack escaping was very low (only 1% occurrence). The implications of this research suggest that the primary management consideration for reducing wild dog escapes from fenced reserves should be maintaining adequate reserve-wide fence integrity, rather than focusing on social structure or drivers of inter- and intra-specific competition.     

Population Genetic Patterns of Threatened European Mudminnow (Umbra krameri Walbaum, 1792) in a Fragmented Landscape: Implications for Conservation Management

The European mudminnow (Umbra krameri) is a Middle Danubian endemic fish species, which is characterised by isolated populations living mainly in artificial habitats in the centre of its range, in the Carpathian Basin. For their long term preservation, reliable information is needed about the structure of stocks and the level of isolation. The recent distribution pattern, and the population genetic structure within and among regions were investigated to designate the Evolutionary Significant, Conservation and Management Units (ESUs, CUs, MUs) and to explore the conservation biological value of the shrinking populations. In total, eight microsatellite loci were studied in 404 specimens originating from eight regions. The results revealed a pronounced population structure, where strictly limited gene flow was detected among regions, as well as various strengths of connections within regions. Following the results of hierarchical structure analyses, two ESUs were supposed in the Carpathian Basin, corresponding to the Danube and Tisza catchments. Our results recommend designating the borders of CUs in an 80–90km range and 16 clusters should be set up as MUs for the 33 investigated populations. How these genetic findings can be used to better allocate conservation resources for the long term maintenance of the metapopulation structure of this threathened endemic fish is discussed.     

Cryptic genetic diversity of <Emphasis Type="Italic">Neverita didyma</Emphasis> in the coast of China revealed by phylogeographic analysis: implications for management and conservation

Revealing cryptic biodiversity and understanding the processes that promote lineage diversification will provide valuable insights into management and protection of exploitable species. Neverita didyma is one of the most common marine species along the coast of China and possesses highly economic and nutritional value. Despite being heavily harvested each year, the genetic diversity of this species has never been assessed in the coastal areas of China. Here, we analyzed the diversity of this species based on the barcode region of the mitochondrial gene Cytochrome Oxidase subunit I (COI) and utilized different species delineation approaches to infer evolutionarily significant units (ESUs). Three distinct ESUs, with high genetic distance among each, were identified. Divergence time estimates suggested that the high genetic distances were probably associated with historical isolation of the marginal seas during Pleistocene low sea level periods. The three ESUs did not map to distinct geographical distribution, possibly attributing to the repeated isolation in different refugia and random postglacial recolonization. Moreover, N. didyma in Haizhou Bay deserves priority protection due to its unique ESU. To improve management regulations in the marine realm, our research also stresses the need for more empirical studies on genetic diversity of commercially exploited species in coastal environments of China.     

Insights into the Management of Large Carnivores for Profitable Wildlife-Based Land Uses in African Savannas

Large African predators, especially lions (Panthera leo) and leopards (Panthera pardus), are financially valuable for ecotourism and trophy hunting operations on land also utilized for the production of other wildlife species for the same purpose. Predation of ungulates used for trophy hunting can create conflict with landholders and trade off thus exists between the value of lions and leopards and their impact on ungulate populations. Therefore productionist and conservation trade-offs are complexly graded and difficult to resolve. We investigated this with a risk-benefit analysis on a large private wildlife production area in Zimbabwe. Our model showed that lions result in substantial financial costs through predation on wild ungulates that may not be offset by profits from hunting them, whereas the returns from trophy hunting of leopards are projected to exceed the costs due to leopard predation. In the absence of additional income derived from photo-tourism the number of lions may need to be managed to minimize their impact. Lions drive important ecological processes, but there is a need to balance ecological and financial imperatives on wildlife ranches, community wildlife lands and other categories of multiple use land used for wildlife production. This will ensure the competitiveness of wildlife based land uses relative to alternatives. Our findings may thus be limited to conservancies, community land-use areas and commercial game ranches, which are expansive in Africa, and should not necessarily applied to areas where biodiversity conservation is the primary objective, even if hunting is allowed there.     

Achieving True Sustainability of Zoo Populations

For the last 30 years, cooperative management of irreplaceable animal populations in zoos and aquariums has focused primarily on the goal of minimizing genetic decay within defined time frames, and large advances have been made in technologies to optimize genetic management of closed populations. However, recent analyses have shown that most zoo programs are not projected to meet their stated goals. This has been described as a lack of achieving “sustainability” of the populations, yet by definition a goal of managed decay is not a plan for sustainability. True sustainability requires management of the resource in manner that does not deplete its value for the future. Achieving such sustainability for many managed populations may require changing from managing isolated populations to managing populations that are part of a broader metapopulation, with carefully considered exchange between populations across a spectrum of ex situ to in situ. Managing zoo populations as components of comprehensive conservation strategies for the species will require research on determinants of various kinds of genetic, physiological, behavioral, and morphological variation and their roles in population viability, development of an array of management techniques and tools, training of population managers in metapopulation management and integrated conservation planning, and projections of impacts of management strategies on the viability of the captive populations and all populations that are interactively managed or affected. Such a shift in goals and methods would result in zoo population management being an ongoing part of species conservation rather than short‐term or isolated from species conservation. Zoo Biol. 32:19‐26, 2013. © 2012 Wiley Periodicals, Inc.     

Comparative phylogeography and the identification of genetically divergent areas for conservation

Genetic diversity is recognized as a fundamental component of biodiversity and its protection is incorporated in several conventions and policies. However, neither the concepts nor the methods for assessing conservation value of the spatial distribution of genetic diversity have been resolved. Comparative phylogeography can identify suites of species that have a common history of vicariance. In this study we explore the strengths and limitations of Faith's measure of 'Phylogenetic Diversity' (PD) as a method for predicting from multiple intraspecific phylogeographies the underlying feature diversity represented by combinations of areas. An advantage of the PD approach is that information on the spatial distribution of genetic diversity can be combined across species and expressed in a form that allows direct comparison with patterns of species distributions. It also seeks to estimate the same parameter, feature diversity, regardless of the level of biological organization. We extend the PD approach by using Venn diagrams to identify the components of PD, including those unique to or shared among areas and those which represent homoplasy on an area tree or which are shared across all areas. PD estimation should be complemented by analysis of these components and inspection of the contributing phylogeographies. We illustrate the application of the approach using mtDNA phylogeographies from vertebrates resident in the wet tropical rainforests of north-east Queensland and compare the results to biodiversity assessments based on the distribution of endemic vertebrate species. The genetic vs. species approaches produce different assessments of conservation value, perhaps reflecting differences in the temporal and spatial scale of the determining processes. The two approaches should be seen as complementary and, in this case, conservation planning should incorporate information on both dimensions of biodiversity.     

Territoriality and spatial patterns of white rhinoceros in Matobo National Park, Zimbabwe

Boundaries associated with reserves or conservation areas may alter spatial patterns of individuals, and may affect the sexes differently in species that exhibit sex-specific patterns of space use. Because of poaching threats, most African rhinos today persist within sanctuaries or reserves with fenced perimeters. We examined spatial patterns of adult white rhinos ( Ceratotherium simum ) in Matobo National Park, Zimbabwe. Matobo Park contains a high-density population of rhinos within a fenced reserve and a lower-density group outside of the fenced area. Adult females in the lower-density group used significantly larger home ranges than those in the high-density group. Overlap among female home ranges was extensive in both groups. Adult male white rhinos establish territories that are exclusive of other males performing scent-marking behaviours, and in Matobo Park, male territories were much larger than those observed in other populations of white rhinos. Additionally, few adult males in the high-density reserve held territories that occupied most available space; two-thirds of adult males were excluded from establishing territories, and followed a non-territorial tactic. In the absence of dispersal opportunities, managers may need to regulate the number of males within smaller reserves. Monitoring of spatial patterns should be undertaken to permit management decisions to be made with an understanding of male behaviour and territorial status.