ABSTRACT Aim Indigenous range maps are fundamental documents in biogeography, phylogeny and conservation. We define the indigenous range of a species as ecoregions (or parts of ecoregions) where the species was likely found before humans became a major factor shaping the species' distribution, beginning at a time when the geographical alignment of the continents and the prevailing climate are (or at least were) roughly consistent with current conditions. We developed a structured, generally applicable method to map a species' indigenous range and applied this process to the tiger ( Panthera tigris ). Location Terrestrial Asia. Methods To guide our mapping, we synthesised a database of over 70,000 tiger observations with dates and locations. We developed a structured Delphi process to assign categories of indigenous range to ecoregions aided by a climate niche model. We analysed tiger habitat change at the ecoregional scale using the anthropogenically modified biomes (‘Anthrome 12K’) dataset to suggest dates of first significant human impact. Finally, we estimated extirpation dates for ecoregions where tigers have been extirpated. Results We found the tiger once occupied a likely indigenous resident range of approximately 11.5 million km 2 , crossing 116 ecoregions. We also mapped an additional c. 11.7 million km 2 of exploratory range and 1.2 million km 2 of possible resident range. Collectively these areas overlap with 36 modern countries. Significant human disruption of the species' habitat seems to have begun over 6000 years ago in some areas, but in other regions has yet to materialise. In few arid ecoregions, human activities appear to have modestly increased habitat availability in the past, yet overall tigers have lost between 90% and 95% of their indigenous range over the last 8500 years. Main Conclusions We define the ‘indigenous range’ of a species, develop a replicable biogeographical procedure, apply the procedure to the tiger and discuss transferability to other species.
Of all the ways human beings have modified the planet over the last 10,000 years, habitat loss is the most important for other species. To address this most critical threat to biodiversity, governments, non-governmental actors, and the public need to know, in near real-time, where and when habitat loss is occurring. Here we present an integrated habitat modelling system at the range-wide scale for the tiger (Panthera tigris) to measure and monitor changes in tiger habitat at range-wide, national, biome, and landscape scales, as often as the underlying inputs change. We find that after nearly 150 years of decline, effective potential habitat for the tiger seems to have stabilized at around 16% of its indigenous extent (1.817 million km2). As of the 1st of January 2020, there were 63 Tiger Conservation Landscapes in the world, covering 911,920 km2 shared across ten of the 30 modern countries which once harbored tiger populations. Over the last 20 years, the total area of Tiger Conservation Landscapes (TCLs) declined from 1.025 million km2 in 2001, a range-wide loss of 11%, with the greatest losses in Southeast Asia and southern China. Meanwhile, we documented expansions of modelled TCL area in India, Nepal, Bhutan, northern China, and southeastern Russia. We find significant potential for restoring tigers to existing habitats, identified here in 226 Restoration Landscapes. If these habitats had sufficient prey and were tigers able to find them, the occupied land base for tigers might increase by 50%. Our analytical system, incorporating Earth observations, in situ biological data, and a conservation-oriented modelling framework, provides the information the countries need to protect tigers and enhance habitat, including dynamic, spatially explicit maps and results, updated as often as the underlying data change. Our work builds on nearly 30 years of tiger conservation research and provides an accessible way for countries to measure progress and report outcomes. This work serves as a model for objective, range-wide, habitat monitoring as countries work to achieve the goals laid out in the Sustainable Development Goals, the 30×30 Agenda, and the Kunming-Montreal Global Biodiversity Framework.
Aim The increasing availability of animal tracking datasets collected across many sites provides new opportunities to move beyond local assessments to enable detailed and consistent habitat mapping at biogeographical scales. However, integrating wildlife datasets across large areas and study sites is challenging, as species' varying responses to different environmental contexts must be reconciled. Here, we compare approaches for large-area habitat mapping and assess available habitat for a recolonizing large carnivore, the Eurasian lynx (Lynx lynx).LocationEurope.Methods We use a continental-scale animal tracking database (450 individuals from 14 study sites) to systematically assess modelling approaches, comparing (1) global strategies that pool all data for training versus building local, site-specific models and combining them, (2) different approaches for incorporating regional variation in habitat selection and (3) different modelling algorithms, testing nonlinear mixed effects models as well as machine-learning algorithms.Results Testing models on training sites and simulating model transfers, global and local modelling strategies achieved overall similar predictive performance. Model performance was the highest using flexible machine-learning algorithms and when incorporating variation in habitat selection as a function of environmental variation. Our best-performing model used a weighted combination of local, site-specific habitat models. Our habitat maps identified large areas of suitable, but currently unoccupied lynx habitat, with many of the most suitable unoccupied areas located in regions that could foster connectivity between currently isolated populations.Main Conclusions We demonstrate that global and local modelling strategies can achieve robust habitat models at the continental scale and that considering regional variation in habitat selection improves broad-scale habitat mapping. More generally, we highlight the promise of large wildlife tracking databases for large-area habitat mapping. Our maps provide the first high-resolution, yet continental assessment of lynx habitat across Europe, providing a consistent basis for conservation planning for restoring the species within its former range.
Context Adjustments in habitat use by large carnivores can be a key factor facilitating their coexistence with people in shared landscapes. Landscape composition might be a key factor determining how large carnivores can adapt to occurring alongside humans, yet broad-scale analyses investigating adjustments of habitat use across large gradients of human pressure and landscape composition are lacking. Objectives Here, we investigate adjustments in habitat use by Eurasian lynx ( Lynx lynx ) in response to varying availability of refuge habitats (i.e., forests and rugged terrain) and human landscape modification. Methods Using a large tracking dataset including 434 individuals from seven populations, we assess functional responses in lynx habitat use across two spatial scales, testing for variation by sex, daytime, and season. Results We found that lynx use refuge habitats more intensively with increasing landscape modification across spatial scales, selecting forests most strongly in otherwise open landscapes and rugged terrain in mountainous regions. Moreover, higher forest availability enabled lynx to place their home ranges in more human-modified landscapes. Human pressure and refuge habitat availability also shaped temporal patterns of lynx habitat use, with lynx increasing refuge habitat use and reducing their use of human-modified areas during periods of high exposure (daytime) or high vulnerability (postnatal period) to human pressure. Conclusions Our findings suggest a remarkable adaptive capacity of lynx towards human pressure and underline the importance of refuge habitats across scales for enabling coexistence between large carnivores and people. More broadly, we highlight that the composition of landscapes determines how large carnivores can adapt to human pressure and thus play an important role shaping large carnivore habitat use and distributions.
Abstract The ecology and evolution of reproductive timing and synchrony have been a topic of great interest in evolutionary ecology for decades. Originally motivated by questions related to behavioral and reproductive adaptation to environmental conditions, the topic has acquired new relevance in the face of climate change. However, there has been relatively little research on reproductive phenology in mammalian carnivores. The Eurasian lynx (Lynx lynx) occurs across the Eurasian continent, covering three of the four main climate regions of the world. Thus, their distribution includes a large variation in climatic conditions, making it an ideal species to explore reproductive phenology. Here, we used data on multiple reproductive events from 169 lynx females across Europe. Mean birth date was May 28 (April 23 to July 1), but was ~10 days later in northern Europe than in central and southern Europe. Birth dates were relatively synchronized across Europe, but more so in the north than in the south. Timing of birth was delayed by colder May temperatures. Severe and cold weather may affect neonatal survival via hypothermia and avoiding inclement weather early in the season may select against early births, especially at northern latitudes. Overall, only about half of the kittens born survived until onset of winter but whether kittens were born relatively late or early did not affect kitten survival. Lynx are strict seasonal breeders but still show a degree of flexibility to adapt the timing of birth to surrounding environmental conditions. We argue that lynx give birth later when exposed to colder spring temperatures and have more synchronized births when the window of favorable conditions for raising kittens is shorter. This suggests that lynx are well adapted to different environmental conditions, from dry and warm climates to alpine, boreal, and arctic climates. This variation in reproductive timing will be favorable in times of climate change, as organisms with high plasticity are more likely to adjust to new environmental conditions.
Taxonomy and systematics are fundamental to the success of conservation actions. A robust and accurate classification of living organisms is vital for understanding biodiversity, using limited resources wisely, prioritising conservation action, and for legal protection and regulation of trade. However, all too often current taxonomies are not based on the latest science and reflect traditional classifications developed in the nineteenth and early twentieth centuries. Understanding of the numbers of species has also changed dramatically with the widespread, but patchy, adoption of the phylogenetic species concept for many vertebrate groups. This has led to a situation where taxonomies are constantly changing in the light of new, mostly genetic, studies. Therefore, the need for a global list of accepted taxa has been recognised by the conservation community as a way of overcoming the uncertainties caused by this dynamic situation. Here, we propose a traffic-light system that may assist the efforts towards the global list of accepted taxa. The traffic-light system indicates the level of certainty in support of the recognition of each taxon, which mainly comprises morphological, genetic and biogeographical lines of evidence. So far, this approach has been adopted by the IUCN Cat Specialist Group to revise felid taxonomy, and the resulting classification has been adopted by the IUCN Red List of Threatened Species and CITES. We discuss the applicability of the approach to other species groups.
Reintroductions may produce populations that suffer from decreasing genetic diversity due to isolation, genetic drift and inbreeding if not assisted by careful management. To assess the genetic outcomes of reintroductions in large carnivores, we used the Eurasian lynx (Lynx lynx) as a case study, which was the subject of several reintroduction attempts over the last 50 years. Although some restocking actions initially appeared successful, lynx recovery has stagnated in recent years. To reveal potential genetic causes of slow lynx recovery in Europe, we examined genome-wide patterns of genetic diversity and inbreeding using single nucleotide polymorphisms (SNPs) in all six successfully reintroduced populations in central Europe, as well as twelve natural populations across Europe and Asia. All reintroduced populations showed lower genetic diversity and elevated levels of inbreeding compared to source and other natural populations. Recent inbreeding is prevalent in all reintroduced populations with varying degrees of severity; the most severe cases are those with the lowest number of founding individuals. Interestingly, we found evidence of lower genetic diversity and recent inbreeding in the source population for five reintroduced populations, begging the question if individuals taken from these source populations can safeguard sufficient genetic diversity for future reintroductions. Given the observed genetic consequences, we advocate for standardized regular genomic assessment of source and target populations as well as individuals prior to release. Our study provides compelling evidence for the serious consequences of founder population size on the genetic diversity of reintroduced large carnivore populations, which has broad implications for their conservation.
Habitat selection is a multi-scale process driven by trade-offs between benefits, such as resource abundance, and disadvantages, such as the avoidance of risk. The latter includes human disturbances, to which large carnivores, with their large spatial requirements, are especially sensitive. We investigated the ecological processes underlying multi-scale habitat selection of a large carnivore, namely Eurasian lynx, across European landscapes characterized by different levels of human modification. Using a unique dataset of 125 lynx from 9 study sites across Europe, we compared used and available locations within landscape and home-range scales using a novel Mixed Effect randomForest approach, while considering environmental predictors as proxies for human disturbances and environmental resources. At the landscape scale, lynx avoided roads and human settlements, while at the home-range scale natural landscape features associated with shelter and prey abundance were more important. The results showed sex was of relatively low variable importance for lynx's general habitat selection behaviour. We found increasingly homogeneous responses across study sites with finer selection scales, suggesting that study site differences determined coarse selection, while utilization of resources at the finer selection scale was broadly universal. Thereby describing lynx's requirement, if not preference, for heterogeneous forests and shelter from human disturbances and implying that regional differences in coarse-scale selection are driven by availability rather than preference. These results provide crucial information for conserving this species in human-dominated landscapes, as well as for the first time, to our knowledge, generalising habitat selection behaviour of a large carnivore species at a continental scale.
AbstractWhen wild‐caught Eurasian lynx (Lynx lynx) from the Slovak Carpathian Mountains were reintroduced to Central Switzerland in the early 1970s and spread through the north‐western Swiss Alps (NWA), they faced a largely unfamiliar landscape with strongly fragmented forests, high elevations, and intense human land use. For more than 30 years, radio‐collared lynx have been monitored during three different project periods (in the 1980s, 1990s, and 2010s). Our study explored, how lynx over generations have learned to adjust to the alpine environment. We predicted that (1) lynx nowadays select more strongly for open habitats, higher elevations, and steep slopes compared to the early stages of recolonization and that (2) consequently, there were significant changes in the Eurasian lynx’ prey spectrum. To test our predictions, we analyzed telemetry data (VHF, GPS) of 13 adult resident lynx in the NWA over 35 years, using Resource Selection Functions. Furthermore, we compared kills recorded from different individuals inhabiting the same region during three project periods. In general, lynx preferred forested areas, but over the years, they avoided open habitat less. Compared to the early stage of the recolonization, lynx in the most recent project period selected for higher elevations and the proportion of chamois in their prey spectrum surmounted that of roe deer. Potential driving factors for the observed changes could be increasing tolerance to human presence, intraspecific competition, or fitness benefits through exploitation of new resources. Long‐term studies like ours provide important insight into how animals can respond to sudden environmental changes, e.g., in the course of translocations into new areas or anthropogenic alterations of their habitats.
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Switzerland has become an important source of Eurasian lynx Lynx lynx for reintroduction projects in Europe. It is now widely accepted that translocations of animals are associated with a serious health risk. Therefore, the development of multidisciplinary expertise and the elaboration of veterinary protocols are needed, which require knowledge on the health status of the source population and information on potential health risks at the release site. Here, both disease cases and carriers of potentially threatening pathogens have to be taken into consideration. In Switzerland, a range of infectious agents circulate within lynx populations apparently without associated morbidity. However, genetic analyses combined with health investigations have pointed at a possible inbreeding depression. Furthermore, unexpected health issues arose in the framework of translocations. Overall, the Swiss experiences underline the necessity of long-term health surveillance of reintroduced and small isolated wildlife populations, the usefulness of well-established veterinary protocols in the framework of translocation projects, the value of multidisciplinary collaborations and of sample archives for further analyses, and the need for adaptive management based on scientific data. For a conservation programme of the Eurasian lynx on a pan-European level, procedure harmonisation should be sought.
The Eurasian lynx (Lynx lynx) population in Switzerland serves as a source for reintroductions in neighboring countries. In 2016–2017, three lynx from the same geographical area were found seropositive for feline immunodeficiency virus (FIV) in the framework of an international translocation program. This novel finding raised questions about the virus origin and pathogenicity to lynx, the emerging character of the infection, and the interpretation of serological results in other lynx caught for translocation. Archived serum samples from 84 lynx captured in 2001–2016 were retrospectively tested for FIV antibodies by Western blot. All archived samples were FIV-negative. The three seropositive lynx were monitored in quarantine enclosures prior to euthanasia and necropsy. They showed disease signs, pathological findings, and occurrence of co-infections reminding of those described in FIV-infected domestic cats. All attempts to isolate and characterize the virus failed but serological data and spatiotemporal proximity of the cases suggested emergence of a lentivirus with antigenic and pathogenic similarities to FIV in the Swiss lynx population. A decision scheme was developed to minimize potential health risks posed by FIV infection, both in the recipient and source lynx populations, considering conservation goals, animal welfare, and the limited action range resulting from local human conflicts. Development and implementation of a cautious decision scheme was particularly challenging because FIV pathogenic potential in lynx was unclear, negative FIV serological results obtained within the first weeks after infection are unpredictable, and neither euthanasia nor repatriation of multiple lynx was acceptable options. The proposed scheme distinguished between three scenarios: release at the capture site, translocation, or euthanasia. Until April 2021, none of the 40 lynx newly captured in Switzerland tested FIV-seropositive. Altogether, seropositivity to FIV was documented in none of 124 lynx tested at their first capture, but three of them seroconverted in 2016–2017. Diagnosis of FIV infection in the three seropositive lynx remains uncertain, but clinical observations and pathological findings confirmed that euthanasia was appropriate. Our experiences underline the necessity to include FIV in pathogen screenings of free-ranging European wild felids, the importance of lynx health monitoring, and the usefulness of health protocols in wildlife translocation.
Abstract With an estimated < 50 adult individuals remaining, the Critically Endangered Balkan lynx Lynx lynx balcanicus is one of the rarest, most threatened and least-studied large carnivores. To identify priority conservation areas and actions for the subspecies, during 2006–2014 we conducted 1,374 questionnaire surveys throughout the potential range of the Balkan lynx to (1) evaluate human–lynx interactions and identify potential threats, and (2) determine the probability of site use in 207 grid cells through occupancy modelling. Human–lynx interactions were related mainly to poaching of lynx, and damage to livestock by lynx. Poaching was intense throughout the potential range of the subspecies, apparently having affected 50–100% of the total estimated extant population. Damage to livestock was recorded only in relation to sheep, mainly in the southern part of the lynx's potential range. Occupancy modelling indicated 108 grid cells with high probability of site use, which was affected mainly by increased terrain ruggedness and reduced forest cover. Based on the combined results of our study we identified five priority areas for conservation, as well as in situ habitat protection, community participation in the conservation of the subspecies, and the improvement and implementation of the existing legal framework as the priority conservation actions for the Balkan lynx.
The Eurasian lynx Lynx lynx population in the Carpathian Mountains is considered to be one of the best preserved and largest in Europe and hence is a source for past and current reintroduction projects in central Europe. However, its status in Slovakia has been reported to the European Commission on the basis of hunters ' reports and expert estimates that have never been validated by a robust scientific approach. We conducted the first camera-trapping surveys to estimate the density of Eurasian lynx in Slovakia by means of spatial capture-recapture models in two reference areas during 2011-2015. We estimated population density per 100 km(2) of suitable lynx habitat (posterior SD) as 0.58 +/- SD 0.13 independent individuals (adults and subadults) in the Stiavnica Mountains and 0.81 +/- SD 0.29 in Veka Fatra National Park and surroundings. These are the lowest densities estimated using spatial capture-recapture models so far reported for the species, suggesting the lynx population in Slovakia is below carrying capacity. We suspect that low densities may be attributable to undetected human-caused mortality. Our results imply that official game statistics are substantially overestimated. Moreover, the lynx population in Slovakia may not be at favourable conservation status as required by the EU Habitats Directive. We therefore call for a thorough assessment of the density and trend of the Slovak Carpathian lynx population, and the establishment of a scientifically robust monitoring system.
The Eurasian lynx is of special conservation concern based on the European Union's Habitat Directive and its populations need to be maintained or restored at favourable conservation status. To evaluate lynx population status, appropriate monitoring needs to be in place. We modelled the distribution dynamics of lynx in the Alps (200 000 km 2 ) during 1994–2014 at a resolution of 100 km 2 . Lynx distribution and detection probability varied by year, country, forest cover, elevation and distance to the nearest release site. Occupancy of neighbouring quadrats had a strong positive effect on colonization and persistence rates. Our analyses demonstrate the importance of accounting for imperfect detection: the raw data underestimated the lynx range by 55% on average, depending on country and winter. Over the past 20 years the Alpine lynx range has expanded at an average rate of 4% per year, which was partly due to the lynx translocations to new areas. Our approach to large‐scale distribution modelling and analysing trends using site occupancy models can be applied retrospectively and is useful in many cases where a network of trained people is established to report the presence of target species, for example, in Europe where member states of the European Union have to report conservation status of species of community interest. Hence, dynamic occupancy models are an appealing framework for inference about the large‐scale range dynamics based on opportunistic data and a useful tool for large‐scale management and conservation programmes.
The leopard cat Prionailurus bengalensis is one of the smallest cat species found on Borneo and is the most widely distributed wild cat species in Asia. It is listed on The IUCN Red List of Threatened Species as Least Concern. The leopard cat is known to tolerate habitat disturbance and to occur in a range of vegetation types including primary and secondary forests, plantations and orchards. However, the extent to which the leopard cat can tolerate habitat disturbance and utilise non-forest areas is still unknown. In this paper, we collected 373 leopard cat location records across Borneo; of these, 228 records were obtained between 2001 and 2011. We analysed 63 (Balanced Model) and 102 (Spatial Filtering Model) records to model habitat suitability. The models predicted more than two-thirds of Borneo to be suitable habitat for the leopard cat. Almost the entire area of Sabah, Sarawak, Brunei and East Kalimantan were predicted to be suitable. The south of Central Kalimantan and the majority of West Kalimantan were predicted to be marginally suitable whereas half of South Kalimantan was predicted to be unsuitable for this species. Although more intensive surveys in Brunei, West Kalimantan and South Kalimantan would help to understand leopard cat distribution, no conservation actions are needed for this species.
The leopard’s ( Panthera pardus ) broad geographic range, remarkable adaptability, and secretive nature have contributed to a misconception that this species might not be severely threatened across its range. We find that not only are several subspecies and regional populations critically endangered but also the overall range loss is greater than the average for terrestrial large carnivores. To assess the leopard’s status, we compile 6,000 records at 2,500 locations from over 1,300 sources on its historic (post 1750) and current distribution. We map the species across Africa and Asia, delineating areas where the species is confirmed present, is possibly present, is possibly extinct or is almost certainly extinct. The leopard now occupies 25–37% of its historic range, but this obscures important differences between subspecies. Of the nine recognized subspecies, three ( P. p. pardus, fusca, and saxicolor ) account for 97% of the leopard’s extant range while another three ( P. p. orientalis, nimr, and japonensis ) have each lost as much as 98% of their historic range. Isolation, small patch sizes, and few remaining patches further threaten the six subspecies that each have less than 100,000 km 2 of extant range. Approximately 17% of extant leopard range is protected, although some endangered subspecies have far less. We found that while leopard research was increasing, research effort was primarily on the subspecies with the most remaining range whereas subspecies that are most in need of urgent attention were neglected.
Little is known about the ecology of the rare marbled cat Pardofelis marmorata on Borneo. In addition, the little information that is available on the species often comes from incidental sightings. Here we use the MaxEnt algorithm to produce a habitat suitability map for this species based on a compilation of existing data. We collected 105 marbled cat occurrence records for Borneo, of which 27 (Balanced Model) or 69 (Spatial Filtering Model) were used to estimate potential habitat suitability. The resulting relative habitat suitability map showed key conservation areas in Borneo. According to these results it appears that the most suitable habitats for marbled cat are lowland forests, but these forests are most threatened by deforestation and other anthropogenic activities. It is imperative to develop appropriate conservation strategies for the marbled cat on Borneo, including long-term research and monitoring, reduction of human disturbances in lowland forests, increased data-sharing and research networking, and stakeholder involvement for conservation planning and activities.