Animal behavior informed by history: Was the Asiatic cheetah an obligate gazelle hunter?

Understanding key ecological adaptations, such as foraging, when a predator is almost extinct is complex. Nonetheless, that information is vital for the recovery of the persisting individuals. Therefore, reviewing historical, ethnobiological and recent records can assist in exploring the species behavioral ecology. We applied this approach to Asiatic cheetahs (Acinonyx jubatus venaticus), which once roamed most west and central Asian countries but now is confined to a few dozens in Iran, at historical (pre-1970) and recent (post-1970) scales. We addressed a widely popular perception that Asiatic cheetahs were subjected to prey shifts from gazelles (Gazella spp.) in open plains areas to urial (Ovis vignei) in mountains because of gazelle populations declines due to anthropogenic influences. We also quantified recent prey choice of Asiatic cheetahs and their behavioral plasticity in foraging different prey species types. Although ethnobiological and historical records suggested that gazelle species were the main prey for cheetahs across their Asian range. However, urial were also commonly reported to be hunted by cheetahs across their historical Asian range, showing that the predation on mountain ungulates is not an emerging hunting behavior in Asiatic cheetahs. We found spatiotemporal plasticity in recent hunting behavior of cheetahs with selective predation on adult urial males. There was temporal overlap in hunting times for plains dwelling versus mountain ungulates, albeit with some minor differences with morning mostly for gazelles while the predation on mountain ungulates was predominantly post-midday. We provided three management implications for the recovery and restoration of cheetahs in Asia. Our work highlighted the importance of historical studies in informing the behavioral ecology of rare species.

What's on the horizon for community-based conservation? Emerging threats and opportunities.

Community-based conservation can support livelihoods and biodiversity, while reinforcing local and Indigenous values, cultures, and institutions. Its delivery can help address cross-cutting global challenges, such as climate change, conservation, poverty, and food security. Therefore, understanding trends in community-based conservation is pertinent to setting and implementing global goals. We undertook a horizon scan to prioritize 15 emerging threats and opportunities expected to impact the future effectiveness of community-based conservation. Topics relate to global biodiversity policy; human rights; shifting human geography; inclusion, diversity, equity, and access; conservation finance and income; and economic reforms. Our findings offer guidance on strengthening community-based conservation to achieve global environmental and development goals.

Mineral lick use by a community of large herbivores in northern Iran.

Natural mineral licks are ecologically valuable resources to meet the physiological needs of herbivores, particularly in temperate forests. Importantly, licking sites can harbor high anthropogenic risk for conservation-dependent herbivores through higher chance of pathogen spillover from livestock and increased levels of poaching risks. However, to the best of our knowledge, there is no information on the mineral lick use in temperate forests of west Asia and the Caucasus where a few threatened deer species exist. We monitored four naturally occurring mineral licks in Central Alborz Protected Area, northern Iran during May-July 2019 using camera traps and analyzed the mineral content of the licking sites. A total of 53 independent mineral lick visits were obtained from only three species of herbivores, i.e., Caspian red deer (Cervus elaphus maral; n = 21), roe deer (Capreolus capreolus; n = 26), and wild pig (Sus scrofa; n = 6). The sex ratio of visiting Caspian red deer was highly skewed toward females (3M:18F), whereas it was more balanced in visiting roe deer (11M:15F). The species-level distribution of visits corresponded to diurnal and cathemeral for Caspian red deer and for roe deer, respectively, without any evidence of different activity curves. There was a negative nonlinear relationship between the ambient temperature and the visitation rate. Our findings showed that mineral licks are important habitat features for these large herbivores and need to be included in spatial mapping and habitat protection measures.

Current trends suggest most Asian countries are unlikely to meet future biodiversity targets on protected areas.

Aichi Target 11 committed governments to protect ≥17% of their terrestrial environments by 2020, yet it was rarely achieved, raising questions about the post-2020 Global Biodiversity Framework goal to protect 30% by 2030. Asia is a challenging continent for such targets, combining high biodiversity with dense human populations. Here, we evaluated achievements in Asia against Aichi Target 11. We found that Asia was the most underperforming continent globally, with just 13.2% of terrestrial protected area (PA) coverage, averaging 14.1 ± SE 1.8% per country in 2020. 73.1% of terrestrial ecoregions had <17% representation and only 7% of PAs even had an assessment of management effectiveness. We found that a higher agricultural land in 2015 was associated with lower PA coverage today. Asian countries also showed a remarkably slow average annual pace of 0.4 ± SE 0.1% increase of PA extent. These combined lines of evidence suggest that the ambitious 2030 targets are unlikely to be achieved in Asia unless the PA coverage to increase 2.4-5.9 times faster. We provided three recommendations to support Asian countries to meet their post-2020 biodiversity targets: complete reporting and the wider adoption "other effective area-based conservation measures"; restoring disturbed landscapes; and bolstering transboundary PAs.

Intraspecific interactions in a high-density leopard population.

Although less studied than interspecific interactions, interactions among members of the same species can influence space use and temporal activity. Using techniques commonly applied to the analysis of interspecific interactions-multispecies occupancy modeling and the analysis of temporal activity patterns-we studied intraspecific interactions within a high-density population of Persian leopards (Panthera pardus saxicolor) in Tandoureh National Park, northeastern Iran. Using camera-trap data, we investigated spatiotemporal interactions between male leopards, lone female leopards, and families (cubs/females with cubs). While we hypothesized that male and female leopards would display different temporal activity patterns, we did not predict spatial avoidance between these groups. We also predicted that leopard families would exhibit spatiotemporal avoidance from male leopards due to the risk of infanticide. Contrary to our expectations, we did not find any evidence for spatial or temporal avoidance between leopard families and adult male leopards. Male and lone female leopards exhibited positive pairwise co-occurrence, consistent with reports of high overlap between male and female leopard home ranges. While a high level of overlap in temporal activity patterns was found between males/lone females and males/families, there was evidence for variation in the proportion of time each leopard group was active in particular periods of the diel cycle. Male leopards showed cathemeral activity, while lone females and families were more active during daylight hours. The application of these techniques to interactions within a species has improved understanding of the ecology and behavior of this endangered solitary carnivore.

Effects of body size on estimation of mammalian area requirements.

Accurately quantifying species' area requirements is a prerequisite for effective area-based conservation. This typically involves collecting tracking data on species of interest and then conducting home-range analyses. Problematically, autocorrelation in tracking data can result in space needs being severely underestimated. Based on the previous work, we hypothesized the magnitude of underestimation varies with body mass, a relationship that could have serious conservation implications. To evaluate this hypothesis for terrestrial mammals, we estimated home-range areas with global positioning system (GPS) locations from 757 individuals across 61 globally distributed mammalian species with body masses ranging from 0.4 to 4000 kg. We then applied block cross-validation to quantify bias in empirical home-range estimates. Area requirements of mammals <10 kg were underestimated by a mean approximately15%, and species weighing approximately100 kg were underestimated by approximately50% on average. Thus, we found area estimation was subject to autocorrelation-induced bias that was worse for large species. Combined with the fact that extinction risk increases as body mass increases, the allometric scaling of bias we observed suggests the most threatened species are also likely to be those with the least accurate home-range estimates. As a correction, we tested whether data thinning or autocorrelation-informed home-range estimation minimized the scaling effect of autocorrelation on area estimates. Data thinning required an approximately93% data loss to achieve statistical independence with 95% confidence and was, therefore, not a viable solution. In contrast, autocorrelation-informed home-range estimation resulted in consistently accurate estimates irrespective of mass. When relating body mass to home range size, we detected that correcting for autocorrelation resulted in a scaling exponent significantly >1, meaning the scaling of the relationship changed substantially at the upper end of the mass spectrum.

Efectos del Tamaño Corporal sobre la Estimación de los Requerimientos de Área de Mamíferos Resumen La cuantificación precisa de los requerimientos de área de una especie es un prerrequisito para que la conservación basada en áreas sea efectiva. Esto comúnmente implica la recolección de datos de rastreo de la especie de interés para después realizar análisis de la distribución local. De manera problemática, la autocorrelación en los datos de rastreo puede resultar en una subestimación grave de las necesidades de espacio. Con base en trabajos previos, formulamos una hipótesis en la que supusimos que la magnitud de la subestimación varía con la masa corporal, una relación que podría tener implicaciones serias para la conservación. Para probar esta hipótesis en mamíferos terrestres, estimamos las áreas de distribución local con las ubicaciones en GPS de 757 individuos de 61 especies de mamíferos distribuidas mundialmente con una masa corporal entre 0.4 y 4,000 kg. Después aplicamos una validación cruzada en bloque para cuantificar el sesgo en estimaciones empíricas de la distribución local. Los requerimientos de área de los mamíferos <10 kg fueron subestimados por una media ∼15% y las especies con una masa ∼100 kg fueron subestimadas en ∼50% en promedio. Por lo tanto, encontramos que la estimación del área estaba sujeta al sesgo inducido por la autocorrelación, el cual era peor para las especies de talla grande. En combinación con el hecho de que el riesgo de extinción incrementa conforme aumenta la masa corporal, el escalamiento alométrico del sesgo que observamos sugiere que la mayoría de las especies amenazadas también tienen la probabilidad de ser aquellas especies con las estimaciones de distribución local menos acertadas. Como corrección, probamos si la reducción de datos o la estimación de la distribución local informada por la autocorrelación minimizan el efecto de escalamiento que tiene la autocorrelación sobre las estimaciones de área. La reducción de datos requirió una pérdida de datos del ∼93% para lograr la independencia estadística con un 95% de confianza y por lo tanto no fue una solución viable. Al contrario, la estimación de la distribución local informada por la autocorrelación resultó en estimaciones constantemente precisas sin importar la masa corporal. Cuando relacionamos la masa corporal con el tamaño de la distribución local, detectamos que la corrección de la autocorrelación resultó en un exponente de escalamiento significativamente >1, lo que significa que el escalamiento de la relación cambió sustancialmente en el extremo superior del espectro de la masa corporal.

Ex situ management as insurance against extinction of mammalian megafauna in an uncertain world.

The persistence of endangered species may depend on the fate of a very small number of individual animals. In situ conservation alone may sometimes be insufficient. In these instances, the International Union for Conservation of Nature provides guidelines for ex situ conservation and the Convention on Biological Diversity (CBD) indicates how ex situ management can support the CBD's objectives by providing insurance policies for species. The circumstances that justify its use are uncertain. To evaluate the current in situ extinction risk and ex situ management of 43 critically endangered species of mammalian megafauna, we used nonmetric multidimensional scaling and geopolitical variables related to governance, economics, and national policy within their extant ranges. We then fitted generalized additive models to assess the contribution of each variable to the ordination. Fifteen (almost one-third) of the world's terrestrial mammalian megafauna are not the subject of any ex situ management. Seventy-three percent of these taxa occur in areas characterized by political uncertainty, such as border zones or areas affected by armed conflicts, mainly in Africa and the Middle East. A further 23% of taxa in ex situ programs do not meet sustainability criteria for inbreeding avoidance. Strategic conservation planning, such as the One Plan approach, may improve ex situ management for these taxa. Given the escalating trend in threats afflicting megafauna, ex situ management should be considered more rigorously, particularly in politically unstable regions, to achieve CBD Target 12 (prevent extinction of threatened species).

Manejo Ex Situ como Protección contra la Extinción de la Megafauna de Mamíferos en un Mundo Incierto Resumen La persistencia de las especies en peligro puede depender del destino de un número muy pequeño de animales individuales. La conservación in situ por sí sola a veces puede ser insuficiente. Bajo estas instancias, la Unión Internacional para la Conservación de la Naturaleza proporciona directrices para la conservación ex situ y la Convención sobre la Diversidad Biológica (CBD) indica cómo el manejo ex situ puede apoyar a lograr sus objetivos al proporcionar políticas de protección para las especies. Las circunstancias que justifican el uso del manejo ex situ son inciertas. Para evaluar el actual riesgo de extinción in situ y el manejo ex situ de 43 especies de megafauna de mamíferos en peligro crítico de extinción usamos un escalamiento multidimensional no métrico, así como variables geopolíticas relacionadas con el gobierno, la economía y las políticas nacionales dentro de la distribución actual de estas especies. Después ajustamos los modelos sumativos generalizados para evaluar la contribución de cada variable a la ordinación. Quince (casi un tercio) de los mamíferos terrestres pertenecientes a la megafauna no están sujetos a ningún tipo de manejo ex situ. El 73% de estos taxones ocurren en áreas conocidas por su incertidumbre política, como zonas fronterizas o áreas afectadas por conflictos armados, principalmente en África y en el Oriente Medio. Además, un 23% de los taxones que sí se encuentran en programas de conservación ex situ no cumplen con los criterios de sustentabilidad para evitar la endogamia. La planeación de la conservación estratégica, como la estrategia de One Plan, podría mejorar el manejo ex situ para estos taxones. Dada la tendencia creciente en las amenazas que afectan a la megafauna, el manejo ex situ debería considerarse de manera más rigurosa, particularmente en las regiones con inestabilidad política, para alcanzar el Objetivo 12 de la CBD (evitar la extinción de especies amenazadas).

Understanding decision making in a food-caching predator using hidden Markov models.

BACKGROUND: Tackling behavioural questions often requires identifying points in space and time where animals make decisions and linking these to environmental variables. State-space modeling is useful for analysing movement trajectories, particularly with hidden Markov models (HMM). Yet importantly, the ontogeny of underlying (unobservable) behavioural states revealed by the HMMs has rarely been verified in the field. METHODS: Using hidden Markov models of individual movement from animal location, biotelemetry, and environmental data, we explored multistate behaviour and the effect of associated intrinsic and extrinsic drivers across life stages. We also decomposed the activity budgets of different movement states at two general and caching phases. The latter - defined as the period following a kill which likely involves the caching of uneaten prey - was subsequently confirmed by field inspections. We applied this method to GPS relocation data of a caching predator, Persian leopard Panthera pardus saxicolor in northeastern Iran. RESULTS: Multistate modeling provided strong evidence for an effect of life stage on the behavioural states and their associated time budget. Although environmental covariates (ambient temperature and diel period) and ecological outcomes (predation) affected behavioural states in non-resident leopards, the response in resident leopards was not clear, except that temporal patterns were consistent with a crepuscular and nocturnal movement pattern. Resident leopards adopt an energetically more costly mobile behaviour for most of their time while non-residents shift their behavioural states from high energetic expenditure states to energetically less costly encamped behaviour for most of their time, which is likely to be a risk avoidance strategy against conspecifics or humans. CONCLUSIONS: This study demonstrates that plasticity in predator behaviour depending on life stage may tackle a trade-off between successful predation and avoiding the risks associated with conspecifics, human presence and maintaining home range. Range residency in territorial predators is energetically demanding and can outweigh the predator's response to intrinsic and extrinsic variables such as thermoregulation or foraging needs. Our approach provides an insight into spatial behavior and decision making of leopards, and other large felids in rugged landscapes through the application of the HMMs in movement ecology.

A paradox of local abundance amidst regional rarity: the value of montane refugia for Persian leopard conservation.

The population densities of leopards vary widely across their global range, influenced by prey availability, intraguild competition and human persecution. In Asia, particularly the Middle East and the Caucasus, they generally occur at the lower extreme of densities recorded for the species. Reliable estimates of population density are important for understanding their ecology and planning their conservation. We used a photographic spatial capture-recapture (SCR) methodology incorporating animal movement to estimate density for the endangered Persian leopard Panthera pardus saxicolor in three montane national parks, northeastern Iran. We combined encounter history data arising from images of bilaterally asymmetrical left- and right-sided pelage patterns using a Bayesian spatial partial identity model accommodating multiple "non-invasive" marks. We also investigated the effect of camera trap placement on detection probability. Surprisingly, considering the subspecies' reported low abundance and density based on previous studies, we found relatively high population densities in the three national parks, varying between 3.10 ± SD 1.84 and 8.86 ± SD 3.60 individuals/100 km2. The number of leopards detected in Tandoureh National Park (30 individuals) was larger than estimated during comparable surveys at any other site in Iran, or indeed globally. Capture and recapture probabilities were higher for camera traps placed near water resources compared with those placed on trails. Our results show the benefits of protecting even relatively small mountainous areas, which accommodated a high density of leopards and provided refugia in a landscape with substantial human activity.

Ecological correlates of the spatial co-occurrence of sympatric mammalian carnivores worldwide.

The composition of local mammalian carnivore communities has far-reaching effects on terrestrial ecosystems worldwide. To better understand how carnivore communities are structured, we analysed camera trap data for 108 087 trap days across 12 countries spanning five continents. We estimate local probabilities of co-occurrence among 768 species pairs from the order Carnivora and evaluate how shared ecological traits correlate with probabilities of co-occurrence. Within individual study areas, species pairs co-occurred more frequently than expected at random. Co-occurrence probabilities were greatest for species pairs that shared ecological traits including similar body size, temporal activity pattern and diet. However, co-occurrence decreased as compared to other species pairs when the pair included a large-bodied carnivore. Our results suggest that a combination of shared traits and top-down regulation by large carnivores shape local carnivore communities globally.

Anchoring and adjusting amidst humans: Ranging behavior of Persian leopards along the Iran-Turkmenistan borderland.

Understanding the space use and movement ecology of apex predators, particularly in mosaic landscapes encompassing different land-uses, is fundamental for formulating effective conservation policy. The top extant big cat in the Middle East and the Caucasus, the Persian leopard Panthera pardus saxicolor, has disappeared from most of its historic range. Its spatial ecology in the areas where it remains is almost unknown. Between September 2014 and May 2017, we collared and monitored six adult leopards (5 males and 1 female) using GPS-satellite Iridium transmitters in Tandoureh National Park (355 km2) along the Iran-Turkmenistan borderland. Using auto-correlated Kernel density estimation based on a continuous-time stochastic process for relocation data, we estimated a mean home range of 103.4 ± SE 51.8 km2 for resident males which is larger than has been observed in other studies of Asian leopards. Most predation events occurred in core areas, averaging 32.4 ± SE 12.7 km2. Although neighboring leopards showed high spatiotemporal overlap, their hunting areas were largely exclusive. Five out of six of leopards spent some time outside the national park, among human communities. Our study suggests that a national park can play an 'anchoring' role for individuals of an apex predator that spend some time in the surrounding human-dominated landscapes. Therefore, we envisage that instead of emphasizing either land sharing or land sparing, a combined approach can secure the viability of resilient large carnivores that are able to coexist with humans in the rugged montane landscapes of west and central Asia.

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

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

Leopard (Panthera pardus) status, distribution, and the research efforts across its range.

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.