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Fitness costs of reproduction are expected when resources are limited. Costs drive the evolution of life‐history strategies and can affect population dynamics if females change their allocation of resources to reproduction. We studied fitness costs of reproduction in mountain ungulates in Alberta, Canada. We monitored two populations of bighorn sheep (Ovis canadensis) for 44 and 30 years, and one of mountain goats (Oreamnos americanus) for 30 years. Both species are highly iteroparous. Heterogeneity in individual reproductive potential makes fitness costs of reproduction difficult to detect and quantify without manipulations. In capital breeders, individual differences can be partly accounted for by considering body mass and other correlates of reproductive potential. Long‐term monitoring can reveal costs that only manifest under stressful conditions such as disease or resource scarcity. Despite individual differences in reproductive potential, we detected fitness costs of reproduction in females. Costs, in terms of mass gain and survival, are almost entirely born by subsequent offspring, as mothers prioritize their own maintenance and survival. Costs are greater for primiparous females, decrease with increasing body mass and increase as resource availability declines, and sons are costlier than daughters. Costs may increase for senescent females that appear to reduce allocation to reproduction. In bighorn sheep, costs mostly involve reduced mass gain and lower survival of subsequent offspring. In mountain goats, costs include reductions in mass gain, subsequent fecundity and juvenile survival. In males, fitness costs derive mostly from attempts to reproduce rather than from siring success and likely depend upon individual competitiveness. In the absence of selective harvests, dominant males may enjoy high fitness and possibly lower costs compared to subordinates. The conservative reproductive tactic of mountain ungulate females likely explains why density dependence mostly involves later primiparity and lower recruitment, but rarely affects adult survival. Future research will seek to better account for heterogeneity in reproductive potential, assess cumulative reproductive costs and investigate the potential effects of fathers on maternal allocation tactics. Fitness costs of reproduction affect the evolution of life histories, morphology and population dynamics of wild species. They are key to the consequences of different forms of exploitation. The authors combine 104 population‐years of monitoring two species of mountain ungulates to show that fitness costs are mostly transferred to subsequent offspring. Résumé Les coûts de la reproduction émergent lorsque les ressources sont limitées et influencent l’évolution des stratégies d’histoire de vie. Si les femelles modifient l’allocation des ressources dans la reproduction pour mitiger ces coûts, ils peuvent aussi affecter la dynamique de population. Nous avons étudié les coûts de la reproduction chez des ongulés en Alberta, Canada. Nous avons suivi deux populations de mouflons d’Amérique (Ovis canadensis) pour 44 et 30 ans, et une de chèvre de montagne (Oreamnos americanus) pour 30 ans. Ces deux espèces sont itéropares. L’hétérogénéité dans le potentiel reproducteur des individus peut masquer les coûts en fitness liés à la reproduction. Ils sont donc plus souvent détectables à l’aide de manipulation expérimentale. Chez les espèces avec reproduction « sur capital », il est possible de contrôler pour les différences individuelles en incluant des variables liées au potentiel reproducteur telles que la masse corporelle. Les suivis à long terme permettent de détecter des coûts qui se manifestent seulement lors de conditions environnementales défavorables. Malgré les différences en potentiel reproducteur, nous avons détecté des coûts chez les femelles. Ces coûts, tels qu’une réduction en croissance ou en survie, sont presque toujours subis par les jeunes nés lors d’épisodes de reproduction subséquents. En effet, les mères priorisent leur propre maintien et leur survie. Les coûts sont supérieurs pour les femelles primipares et dans les conditions défavorables. Ils sont plus faibles pour les femelles plus lourdes et il semble que les fils soient plus coûteux que les filles. Les coûts augmentent aussi chez les femelles sénescentes qui semblent réduire leur allocation dans la reproduction. Chez les mouflons, les coûts impliquent une réduction du gain en masse et une survie plus faible des jeunes nés lors de reproductions subséquentes. Chez les chèvres de montagne, ces coûts sont accompagnés d’une réduction de la fécondité future. Chez les mâles, les coûts sont surtout influencés par l’investissement en temps et en énergie dans le rut plutôt que par le succès d’accouplement. Ces coûts devraient donc surtout dépendre de l’habileté compétitive individuelle. En absence de chasse sélective, les mâles dominants devraient avoir un fort succès reproducteur avec des coûts potentiellement plus faibles comparativement aux subordonnées. La tactique conservative adoptée par les ongulés de montagne femelles peut expliquer pourquoi les effets densité‐dépendant mènent à un retard dans l’âge à la primiparité et à un recrutement plus faible, mais affectent rarement la survie adulte. Nos recherches futures tenteront de mieux quantifier l’hétérogénéité individuelle, évalueront les coûts cumulatifs à la reproduction et investigueront l’effet potentiel des pères sur les tactiques d’allocation maternelles.

Estimating key state variables such as abundance, survival, and recruitment of wild populations, and their interaction with local conditions such as precipitation, is essential for statedependent decision making and management. Estimation of state variables remains challenging, especially in species such as mountain ungulates inhabiting rugged and remote terrains. Camera traps present a technological advance that can mitigate some of the challenges in state estimation and can be used to enhance existing methods. Here, we combined camera traps, individual markings, and GPS tracking to estimate abundance and survival for the threatened Nubian ibex (Capra nubiana) population of the Judean Desert, Israel. Since precipitation is a key determinant for recruitment in desert ungulates, we complemented our estimates with kid-female ratios and regional precipitation records. Between 2016 and 2019, we individually marked 48 ibex of which 38 were also fitted with GPS collars within the Judean Desert. We performed 6 days of camera-trap surveys each year to estimate abundance for the north Judean Desert population using mark-resight in a robust-design framework. Weight-averaged abundance estimates for 2017–2019, respectively, were as follows: N ¯ ^ = 267 (95% CI = 117–418, SE = 51.27), 201 (95% CI = 115–288, SE = 36.85), and 226 (95% CI = 154–299, SE = 36.94) for females, and N ¯ ^ = 291 (95% CI = 154–299, SE = 36.94), 160 (95% CI = 108–212, SE = 25.19), and 283 (95% CI = 181–384, SE = 29.76) for males. By augmenting data from telemetry with camera traps, we estimated survival via known fates for the entire Judean Desert (north, mid, and south populations). Weight-averaged survival estimates for 2016–2019, respectively, were: S ¯ ^ = 0.99 (95% CI = 0.97–1.00, SE = 0.012), 0.90 (95% CI = 0.24–0.99, SE = 0.153), 0.87 (95% CI = 0.62–0.97, SE = 0.081), and 0.88 (95% CI = 0.69–0.96, SE = 0.062) for females, and S ¯ ^ = 0.77 (95% CI = 0.26–0.97, SE = 0.203), 0.64 (95% CI = 0.33–0.86, SE = 0.151), 0.85 (95% CI = 0.60–0.96, SE = 0.087), and 0.88 (95% CI = 0.69–0.96, SE = 0.062) for males. The interannual trends of abundance and survival estimates corresponded with observed kid-female ratios and the regional precipitation records, reflecting a population decline following 2017's drought and an increase following the increased precipitation in 2018. Our findings substantiate the efficacy of using camera traps to estimate Nubian ibex's key state variables, as well as detecting population trends driven by environmental conditions. Despite being costly, the capacity of statistically-robust methods to estimate key state variables is indispensable, and camera traps make similar methods increasingly more feasible in remote and inaccessible populations.

Ecological theory and empirical studies have demonstrated population-level demographic benefits resulting from a diversity of migratory behaviors with important implications for ecology, conservation, and evolution of migratory organisms. Nevertheless, evaluation of migratory portfolios (i.e., the variation in migratory behaviors across space and time among individuals within populations) has received relatively little attention in migratory ungulates, where research has focused largely on the dichotomous behaviors (e.g., resident and migrant) of partially migratory populations. Using GPS data from 361 female bighorn sheep (Ovis canadensis) across 17 (4 restored, 6 augmented, 7 native) populations in Montana and Wyoming, USA, we (1) characterized migratory portfolios based on behavioral and spatial migratory characteristics and (2) evaluated the relative influence of landscape attributes and management histories on migratory diversity. Native populations, which had been extant on the landscape for many generations, had more diverse migratory portfolios, higher behavioral switching rates, reduced seasonal range fidelity, and broad dispersion of individuals across summer and winter ranges. In contrast, restored populations with an abbreviated history on the landscape were largely non-migratory with a narrow portfolio of migratory behaviors, less behavioral switching, higher fidelity to seasonal ranges, and less dispersion on summer and winter ranges. Augmented populations were more variable and contained characteristics of both native and restored populations. Differences in migratory diversity among populations were associated with management histories (e.g., restored, augmented, or native). Landscape characteristics such as the duration and regularity of green-up, human landscape alterations, topography, and snow gradients were not strongly associated with migratory diversity. We suggest a two-pronged approach to restoring migratory portfolios in ungulates that first develops behavior- specific habitat models and then places individuals with known migratory behaviors into unoccupied areas in an effort to bolster migratory portfolios in restored populations, potentially with synergistic benefits associated with variation among individuals and resulting portfolio effects. Management efforts to restore diverse migratory portfolios may increase the abundance, resilience, and long-term viability of ungulate populations.

Estimating the abundance and density of mountain ungulates is difficult because of rugged and remote terrain, high elevations, and rapidly changing weather. Helicopter surveys could overcome these problems, but researchers have seldom applied helicopter-based survey methods at large spatial scales in mountain terrain. We used helicopters to count introduced Himalayan tahr (Hemitragus jemlahicus) at 117 plots, each of 4 km², in New Zealand's Southern Alps during 2016–2019. The sampling frame was 7,844 km² and we located the plots at the vertices of an 8-km grid superimposed over the sampling frame (i.e., a systematic random sampling design). We conducted 3 repeat counts at each plot during summer–autumn. We used the repeat counts to estimate tahr abundance and density, corrected for imperfect detection, using a dynamic N-mixture model for open populations. We estimated the population of tahr in the sampling frame using design-based, finite sampling methods and model-based inference procedures. The mean estimated density of tahr on each plot varied from zero to 31.7 tahr/km². The mean densities of tahr varied among management units, ranging from 0.3 to 10.7 tahr/km², and exceeded specified intervention densities in 6 of the 7 management units. The total design-based estimate of tahr abundance in the sampling frame was 34,500 (95% CI = 27,750–42,900), with a coefficient of variation (CV) of 0.11. The corresponding model-based estimate of total abundance was similar (34,550, 95% CI = 30,250–38,700) but was substantially more precise (CV = 0.06) than the design-based estimate. The precision of the estimates for the individual management units was also better than that of the design-based estimates, with CVs of < 0.20 for all but 1 management unit. Our study provides a repeatable method for sampling mountain ungulates. More generally, robust estimation of abundance and density of mountain ungulates is possible by combining aerial surveys and open population models with an objective, probabilistic sampling design.

Estimating animal abundance has a key role to play in ecology and conservation, but survey methods are always challenged by imperfect detection. Among the techniques applied to deal with this issue, Double Observer (DO) is increasing in popularity due to its cost‐effectiveness. However, the effort of using DO for surveying large territories can be significant. A DO‐based survey method that allows accurate abundance estimations with reduced effort would increase the applicability of the method. This would have positive effects on the conservation of species which are challenging to survey such as mountain ungulates. We used computer simulations based on real data and a field test to assess the reliability of the DO and of a new proposed survey procedure, the Double Observer Adjusted Survey (DOAS). DOAS is based on total block counts adjusted with some DO surveys conducted in a proportion of the total area only. Such DO surveys are then used to estimate detection probability with a mark‐recapture‐derived approach. We found that full DO is much more accurate than simple block counts for abundance estimations. DOAS is a less demanding alternative to full DO and can produce comparable abundance estimates, at the cost of a slightly lower precision. However, in the DOAS overall detectability has to be estimated within a sufficient number of sites (around a quarter of the total) to obtain a higher precision and avoid large overestimations. Practical implications. DO methods could increase the reliability of abundance estimations in mountain ungulates and other gregarious species. Full DO in particular could allow researchers to obtain unbiased estimations with high precision and its usage is therefore suggested instead of block counts in wildlife monitoring. Given the high costs of full DO, the DOAS procedure could be a viable and cost‐effective survey strategy to improve abundance estimates when resources are scarce. Riassunto Le stime di abbondanza degli animali hanno un ruolo cruciale in ecologia e conservazione, ma conteggi accurati sono sempre ostacolati dall'impossibilità di osservare tutti gli individui presenti. Tra i metodi utilizzati per affrontare questo problema, il Doppio Osservatore (DO) sta guadagnando sempre più popolarità grazie ad un ottimo rapporto costi‐benefici. Tuttavia, l'impiego del DO per monitorare zone estese può richiedere un notevole sforzo di campionamento. Pertanto, un protocollo di monitoraggio basato sul DO che mantenga l'accuratezza delle stime, riducendo al contempo lo sforzo richiesto, renderebbe il metodo più pratico e applicabile. Ciò sarebbe particolarmente vantaggioso per la conservazione di specie difficili da monitorare, come gli ungulati di montagna. In questo studio, abbiamo utilizzato simulazioni al computer basate su dati reali e un test sul campo nel Parco Nazionale Gran Paradiso per valutare l'affidabilità del Doppio Osservatore (DO) e di una nuova procedura di monitoraggio: il conteggio corretto con Doppio Osservatore, o Double Obsever Adjusted Survey (DOAS). Il DOAS combina conteggi totali eseguiti tramite il metodo dei block counts con alcune ripetizioni del DO su una porzione dell'area complessiva. Questi rilievi vengono utilizzati per stimare la contattabilità attraverso un approccio derivato dai metodi di marcatura‐ricattura e per correggere i risultati dei conteggi totali. I risultati dello studio evidenziano come i conteggi condotti interamente con il metodo DO forniscono stime di abbondanza decisamente più accurate rispetto ai block counts. Il DOAS rappresenta un'alternativa che richiede meno sforzo di campionamento rispetto al DO, fornendo stime di abbondanza comparabili ma con una precisione leggermente inferiore. Tuttavia, nel DOAS è fondamentale effettuare ripetizioni con DO in un numero sufficiente di siti (circa un quarto del totale) per ottenere una maggiore precisione ed evitare grosse sovrastime. I metodi basati sul Doppio Osservatore hanno il potenziale di migliorare notevolmente l'affidabilità delle stime di abbondanza, sia per gli ungulati di montagna che per altre specie gregarie. In particolare, l'applicazione del DO sull'intera area di studio consentirebbe di ottenere stime altamente accurate e precise, motivo per cui se ne raccomanda l'uso nel monitoraggio della fauna selvatica in alternativa ai censimenti esaustivi (block counts). Tuttavia, considerando i costi elevati associati al DO, il protocollo DOAS rappresenta una soluzione di monitoraggio valida ed economicamente sostenibile, particolarmente utile in contesti di risorse limitate. We here propose a new census framework for mountain ungulates and other gregarious species, the Double Observer Adjusted Survey (DOAS). DOAS is based on total block counts adjusted with some Double Observer surveys to estimate detectability. The DOAS method could produce reliable abundance estimates, lowering the full double observer census efforts.

Robust population surveys and monitoring of ungulates in mountainous areas are hindered by complex terrain and limited access to remote regions. Distance sampling, a widely used method for surveying ungulates in non‐mountainous habitats, faces challenges in such environments due to violations of key assumptions. In this study, we examine the assumptions underlying distance sampling and explore how they can be addressed in mountainous terrain. We then applied distance sampling to estimate the abundance of bharal ( Pseudois nayaur ) on the Tibetan Plateau and compared the results with estimates from the double‐observer survey method, which is commonly used for mountain ungulates. Both methods produced highly comparable estimates—2189 from distance sampling and 2038 from the double‐observer survey—approximately twice as high as the total count (963). Distance sampling yielded a wider confidence interval (95% CI: 1207–3972) than the double‐observer survey (95% CI: 1405–2670). We recommend using distance sampling when human resources are limited or when the closed population assumption is difficult to meet. Otherwise, the double‐observer method is preferable due to its narrower confidence intervals and greater ability to detect population changes. 山地有蹄类的可靠种群调查和监测因地形复杂和偏远地区难以进入而受阻。距离样线法作为非山地地区常用的种群调查方法, 在山地地形中往往难以满足其假设条件。本文讨论了距离采样的关键假设及其在山地地形中的应对方法。随后, 我们应用距离样线法在青藏高原一条山谷中估算了岩羊的数量, 并与常用于山地有蹄类动物的双观察者法调查结果进行了比较。两种方法得出高度相似的估计值 (距离样线法2189只, 双观察者法2038只), 均为直接计数法 (963只) 的两倍。距离样线法得出的置信区间较宽 (95% CI: 1207‐3972), 而双观察者较窄 (95% CI: 1405‐2670) 。建议在人力有限或难以满足种群闭合假设时采用距离采样, 否则双观察者调查因置信区间较窄更适合检测到种群变化。 This study addresses the challenge of accurately estimating populations of mountain‐dwelling ungulates such as the bharal ( Pseudois nayaur ), which are difficult to survey due to rugged terrain and remote locations. Traditional methods like total counts often underestimate true population sizes because animals can be missed. We tested and compared two advanced survey methods—distance sampling and double‐observer surveys—in a valley on the Tibetan Plateau. Distance sampling estimates population size by recording distances to detected groups along transects, but its assumptions are difficult to meet in mountainous areas. The double‐observer method uses two independent teams to improve detection estimates but requires more personnel and careful timing to avoid disturbing animals. Both methods produced similar population estimates—around 2100 bharal—double the number found by total counts (963). Distance sampling had wider confidence intervals, indicating lower precision, while double‐observer surveys were more precise and better at detecting population changes. We adapted distance sampling by dividing the area into watersheds and designing transects to maximize visual coverage. We recommend using double‐observer surveys when a sufficient number of surveyors are available and the population remains closed during surveys. Distance sampling is more appropriate when resources are limited or when the population may move in and out of the survey areas. With appropriate adjustments, both methods can provide reliable estimates to support the conservation of mountain ungulates, which are key prey for predators such as the snow leopard. Distance sampling and double‐observer surveys produced very similar population estimates (2189 and 2038, respectively), both about twice as high as the total count (963). Distance sampling yielded wider confidence intervals (95% CI: 1207–3972) compared to the double‐observer survey (95% CI: 1405–2670). Key assumptions of distance sampling can be met in rugged terrain by dividing the study area into smaller sampling units (e.g., watersheds) and designing transects to maximize visual coverage rather than adhering strictly to straight‐line or random placement. Distance sampling is recommended when human resources are limited or when maintaining a closed population during the survey is difficult. The double‐observer survey is preferable when sufficient personnel are available, as its narrower confidence intervals improve sensitivity to population changes. 本研究针对山地有蹄类动物 (如岩羊 Pseudois nayaur ) 数量准确估算的难题展开, 这类动物因地形复杂和栖息地偏远, 调查十分困难。传统的总数调查方法常因动物未被发现而低估真实数量。我们在青藏高原一条河谷中测试并比较了两种先进的调查方法——距离样线法和双观察者法调查。距离样线法通过沿样线记录观察到的动物群体距离来估算数量, 但其核心假设在山地环境中往往难以满足。双观察者法则由两组独立调查队伍同时进行, 以提高检测率, 但需要更多人力和合理安排间隔时间以避免干扰动物。两种方法得出的数量估计结果相近, 均约为2100只岩羊, 是直接计数法 (963只) 的两倍。距离样线法的置信区间较宽, 表明其精度较低, 而双观察者法调查置信区间较窄, 更适合检测种群变化。研究调整了距离样线法方法以应对山地的复杂地形, 通过将调查区域划分为多个小流域并最大化视野覆盖来保证核心假设的满足。建议在人力充足且种群在调查期间相对闭合时采用双观察者法；在人力有限或难以保证种群封闭时采用距离样线法。总体而言, 经过合理调整, 两种方法均能提供可靠的种群估计, 为保护雪豹等捕食者关键猎物的山地有蹄类动物提供支持。 距离样线法和双观察者法调查得出的种群估计非常接近 (距离样线法2189只, 双观察者法2038只), 均为直接计数法 (963只) 的两倍。距离样线法的置信区间较宽 (95% CI: 1207‐3972), 而双观察者法较窄 (95% CI: 1405‐2670) 。 通过将调查区域划分为较小的采样单元 (如流域), 并设计样线以最大化视觉覆盖, 而非严格遵循直线或随机布置, 可以满足距离样线法在复杂地形中的关键假设。 建议在人力有限或难以保证种群闭合时使用距离样线法；在人力充足且种群相对稳定时优先采用双观察者法调查, 因其置信区间较窄, 更能灵敏地检测种群变化。

Aim Seasonal migrations, such as those of ungulates, are particularly threatened by habitat transformations and fragmentation, climate and other environmental changes caused by anthropogenic activities. Mountain ungulate migrations are neglected because they are relatively short, although traversing heterogeneous altitudinal gradients particularly exposed to anthropogenic threats. Detecting migration routes of these species and understanding their drivers are therefore of primary importance to predict connectivity and preserve ecosystem functions and services. The populations of Alpine ibex Capra ibex have all been reintroduced from the last remnant source population. Despite a general increase in abundance and overall distribution range, ibex populations are mostly disconnected but display intra‐population migrations. Therefore, its conservation is strictly linked to the interplay between external threats and related behavioural responses, including space use and migration. Location Austria, France, Italy and Switzerland. Methods By using 337 migratory tracks from 425 GPS‐collared individuals from 15 Alpine ibex populations distributed across their entire range, we (i) identified the environmental drivers of movement corridors in both spring and autumn and (ii) compared the ability of a connectivity modelling algorithm to predict migratory movements between seasonal ranges of the 15 populations, using either population‐specific or multipopulation datasets, and three validation procedures. Results Steep, south‐facing, snow‐free slopes were selected while high elevation changes were avoided. This revealed the importance of favourable resources and an attempt to limit energy expenditures and perceived predation risk. The abilities of the modelling methods we compared to predict migratory connectivity from the results of those movement analyses were similar. Main Conclusions The trade‐off between energy expenditure, food and cover was the major driver of migration routes and was overall consistent among populations. Based on these findings, we provided useful connectivity models to inform conservation of Alpine ibex and its habitats, and a framework for future research investigating connectivity in migratory species.

Background Predators have the potential to affect prey ecology through both direct effects on population dynamics or indirect effects on behaviour, e.g., by triggering antipredator strategies. Direct effects of predation on single prey species may be limited in ecosystems hosting alternative prey, possibly being overwhelmed by indirect effects. The novel exposure to a predator would provide the opportunity to test for immediate prey responses, but information is scanty for areas recolonised by carnivores. We took advantage of the natural expansion of the wolf Canis lupus in a protected area of western Alps hosting five ungulate species to test the potential for direct versus indirect effects on the main prey, i.e., a widespread mountain herbivore (the Northern chamois Rupicapra rupicapra ). After verifying the contribution of the latter to the diet of the former, we used a semi-experimental (before vs. after) approach by comparing chamois demography, elevation used and group size between two valleys with different recolonisation time (Site A: medium-term vs. Site B: short-term). Results Scat analyses ( N  = 335 samples) indicated that chamois were the staple in the wolf diet in both valleys. Analyses of counts throughout 21 years supported no direct effect of wolf on chamois abundance and survival. Following wolf recolonisation, female chamois ( n  = 3594 observations) in Site A were observed at average elevations 137 m higher compared to the former period, and a concurrent decrease of group size was reported; these effects were not detected in Site B. The same trend was not observed in temperature, precipitation or NDVI, providing no support to a weather- or resource-mediated uplift. Conclusions Although direct/indirect effects of current changes in weather patterns on the observed uplift of chamois may not be ruled out, our results suggest antipredator behaviour as a main determinant of chamois upshift. Finally, we discuss the role of indirect versus direct short-term prey responses in complex ecosystems.

Climate change has led to shifts in species distribution and become a crucial factor in the extinction of species. Increasing average temperatures, temperature extremes, and unpredictable weather events have all become a part of a perfect storm that is threatening ecosystems. Higher altitude habitats are disproportionately affected by climate change, and habitats for already threatened specialist species are shrinking. The Siberian ibex, Capra sibirica, is distributed across Central Asia and Southern Siberia and is the dominant ungulate in the Pamir plateau. To understand how climate change could affect the habitat of Siberian ibex in the Taxkorgan Nature Reserve (TNR), an ensemble species distribution model was built using 109 occurrence points from a four-year field survey. Fifteen environmental variables were used to simulate suitable habitat distribution under different climate change scenarios. Our results demonstrated that a stable, suitable habitat for Siberian ibex was mostly distributed in the northwest and northeast of the TNR. We found that climate change will further reduce the area of suitable habitat for this species. In the scenarios of RCP2.6 to 2070 and RCP8.5 to 2050, habitat loss would exceed 30%. In addition, suitable habitats for Siberian ibex will shift to higher latitudes under climate change. As a result, timely prediction of the distribution of endangered animals is conducive to the conservation of the biodiversity of mountain ecosystems, particularly in arid areas.

The hangul Cervus hanglu hanglu, a Critically Endangered mountain ungulate of Jammu and Kashmir, India, faces the imminent threat of population loss and extinction. Effective management of its largest viable population in Dachigam National Park in the Kashmir Himalaya requires reliable demographic information. Using 14 microsatellite markers we identified 293 individuals (208 females and 85 males) through faecal analysis, and generated data on the genetic status and population size of the hangul in its winter habitat. The mean expected and observed heterozygosities of 0.62 and 0.59 are comparable to those of several red deer Cervus elaphus populations elsewhere. The effective population sizes were 46.3 and 93.7 when the frequencies of rare alleles were considered to be 0.050 and 0.010, respectively. The average mean kinship of the population was 0.34, and there was no evidence of a recent bottleneck event. In genetic mark–recapture analysis the best model included an effect of sex on both detection and recapture probabilities. Detection of males was highest in November, coinciding with the hangul breeding season, whereas detection of females was highest in December. Our estimate of the hangul population using genetic mark–recapture with bootstrapping was 394 individuals. To our knowledge, this is the first study to use genetic data to estimate the population of the hangul. It will guide future studies of this subspecies and also serve as an impetus for identifying founder animals for captive breeding, and for connecting the population in Dachigam National Park with the other small, isolated populations to ensure the long-term survival of this subspecies.