Multi-generational benefits of genetic rescue.

Genetic rescue-an increase in population fitness following the introduction of new alleles-has been proven to ameliorate inbreeding depression in small, isolated populations, yet is rarely applied as a conservation tool. A lingering question regarding genetic rescue in wildlife conservation is how long beneficial effects persist in admixed populations. Using data collected over 40 years from 1192 endangered Florida panthers (Puma concolor coryi) across nine generations, we show that the experimental genetic rescue implemented in 1995-via the release of eight female pumas from Texas-alleviated morphological, genetic, and demographic correlates of inbreeding depression, subsequently preventing extirpation of the population. We present unequivocal evidence, for the first time in any terrestrial vertebrate, that genetic and phenotypic benefits of genetic rescue remain in this population after five generations of admixture, which helped increase panther abundance (> fivefold) and genetic effective population size (> 20-fold). Additionally, even with extensive admixture, microsatellite allele frequencies in the population continue to support the distinctness of Florida panthers from other North American puma populations, including Texas. Although threats including habitat loss, human-wildlife conflict, and infectious diseases are challenges to many imperiled populations, our results suggest genetic rescue can serve as an effective, multi-generational tool for conservation of small, isolated populations facing extinction from inbreeding.

Decline in small mammal species richness in coastal-central California, 1997-2013.

The richness and composition of a small mammal community inhabiting semiarid California oak woodland may be changing in response to climate change, but we know little about the causes or consequence of these changes. We applied a capture-mark-recapture model to 17 years (1997-2013) of live trapping data to estimate species-specific abundances. The big-eared woodrat was the most frequently captured species in the area, contributing 58% of total captures. All small mammal populations exhibited seasonal fluctuations, whereas those of the California mouse, brush mouse, and pinyon mouse declined during the study period. We also applied a multispecies dynamic occupancy model to our small mammal detection history data to estimate species richness, occupancy (ψ), detection (p), local extinction (ϵ), and colonization (γ) probabilities, and to discern factors affecting these parameters. We found that ψ decreased from 0.369 ± 0.088 in 1997 to 0.248 ± 0.054 in 2013; γ was lower during the dry season (May-September) than the wet season (October-April) and was positively influenced by total seasonal rainfall (slope parameter, ß = 0.859 ± 0.371; 95% CI = 0.132-1.587). Mean mammalian species richness decreased from 11.943 ± 0.461 in 1997 to 7.185 ± 0.425 in 2013. With highly variable climatic patterns expected in the future, especially increased frequency and intensity of droughts, it is important to monitor small mammal communities inhabiting threatened California oak woodlands.

Reproductive performance of Peregrine falcons relative to the use of organochlorine pesticides, 1946-2021.

Populations of some fish- and meat-eating birds suffered dramatic declines globally following the introduction of organochlorine pesticides during the late 1940s and 1950s. It has been hypothesised that these population declines during the 1950s-1970s were largely driven by a combination of reproductive failure due to eggshell-thinning, egg breakage and embryonic death attributable to DDT and its metabolites, and to enhanced mortality attributable to the more toxic cyclodiene compounds such as aldrin and dieldrin. Using 75 years (1946-2021) of Peregrine falcon (Falco peregrinus) monitoring data (315 unique nest-sites monitored for 6110 nest-years), we studied the breeding performance of a resident Peregrine population in southern Scotland relative to the spatiotemporal pattern of organochlorine pesticide use. We show that (i) Peregrine breeding success and measures of breeding performance increased substantially following the reduction in, and subsequently a complete ban on, the use of organochlorine pesticides; (ii) improvements in Peregrine breeding performance were more dramatic in southeastern Scotland where agriculture was the predominant land use than in southwestern Scotland where there was less arable and more forested land; (iii) Peregrines nesting closer to the coast generally had higher fledging success (that is, a higher proportion of clutches that produced at least one fledgeling) than those nesting inland farther away from the coast; (iv) low temperatures and excessive rain in May negatively affected Peregrine fledging success; and (v) Peregrine abundance increased in parallel with improvements in reproductive performance following the reduction and then complete ban on the use of organochlorine pesticides in the UK. However, recovery was gradual and occurred over four decades, and rate of recovery varied among measures of reproductive performance (egg, nestling and fledgeling production). Our results suggest that the temporal pattern of organochlorine pesticide use strongly influenced Peregrine reproductive parameters but that the pattern of influence differed regionally. Overall results are consistent with the hypothesis that reproductive failure caused by organochlorine pesticides was an important driver of the decline in the south Scottish Peregrine population, and that improvements in all measures of breeding performance following a reduction and eventual ban on organochlorine use facilitated the observed increase in this population.

Non-exploitative human disturbance provides shelter for prey from predator.

Human activities can influence behaviors of predators and prey, as well as predator-prey interactions. Using camera trap data, we investigated whether or to what extent human activities influenced behaviors of predators (tigers and leopards) and prey (sambar deer, spotted deer, wild boar, and barking deer), and predator-prey interactions in the Barandabhar Corridor Forest (BCF), Chitwan District, Nepal. A multispecies occupancy model revealed that the presence of humans altered the conditional occupancy of both prey and predator species. Specifically, the conditional occupancy probability of prey was substantially higher (ψ = 0.91, CI = 0.89-0.92) when humans were present than when humans were absent (ψ = 0.68, CI = 0.54-0.79). The diel activity pattern of most prey species overlapped strongly with humans, whereas predators were generally more active when humans were absent. Finally, the spatiotemporal overlap analysis revealed that human-prey interactions (i.e., the probability that both humans and prey species being present on the same grid at the same hourly period) was ~3 times higher (10.5%, CI = 10.4%-10.6%) compared to spatiotemporal overlap between humans and predators (3.1%, CI = 3.0%-3.2%). Our findings are consistent with the human shield hypothesis and suggest that ungulate prey species may reduce predation risk by using areas with high human activities.

Lifetime inclusive fitness effects of cooperative polygamy in the acorn woodpecker.

Although over 50 y have passed since W. D. Hamilton articulated kin selection and inclusive fitness as evolutionary explanations for altruistic behavior, quantifying inclusive fitness continues to be challenging. Here, using 30 y of data and two alternative methods, we outline an approach to measure lifetime inclusive fitness effects of cooperative polygamy (mate-sharing or cobreeding) in the cooperatively breeding acorn woodpecker Melanerpes formicivorus. For both sexes, the number of offspring (observed direct fitness) declined while the number of young parented by related cobreeders (observed indirect fitness effect) increased with cobreeding coalition size. Combining these two factors, the observed inclusive fitness effect of cobreeding was greater than breeding singly for males, while the pattern for females depended on whether fitness was age-weighted, as females breeding singly accrued greater fitness at younger ages than cobreeding females. Accounting for the fitness birds would have obtained by breeding singly, however, lifetime inclusive fitness effects declined with coalition size for males, but were greater for females breeding as duos compared to breeding singly, due largely to indirect fitness effects of kin. Our analyses provide a road map for, and demonstrate the importance of, quantifying indirect fitness as a powerful evolutionary force contributing to the costs and benefits of social behaviors.

Does coat colour influence survival? A test in a cyclic population of snowshoe hares.

Some mammal species inhabiting high-latitude biomes have evolved a seasonal moulting pattern that improves camouflage via white coats in winter and brown coats in summer. In many high-latitude and high-altitude areas, the duration and depth of snow cover has been substantially reduced in the last five decades. This reduction in depth and duration of snow cover may create a mismatch between coat colour and colour of the background environment, and potentially reduce the survival rate of species that depend on crypsis. We used long-term (1977-2020) field data and capture-mark-recapture models to test the hypothesis that whiteness of the coat influences winter apparent survival in a cyclic population of snowshoe hares (Lepus americanus) at Kluane, Yukon, Canada. Whiteness of the snowshoe hare coat in autumn declined during this study, and snowshoe hares with a greater proportion of whiteness in their coats in autumn survived better during winter. However, whiteness of the coat in spring did not affect subsequent summer survival. These results are consistent with the hypothesis that the timing of coat colour change in autumn can reduce overwinter survival. Because declines in cyclic snowshoe hare populations are strongly affected by low winter survival, the timing of coat colour change may adversely affect snowshoe hare population dynamics as climate change continues.

An experimental analysis of density dependence in meadow voles: Within-season and delayed effects.

Wild mammal populations exhibit a variety of dynamics, ranging from fairly stable with little change in population size over time to high-amplitude cyclic or erratic fluctuations. A persistent question in population ecology is why populations fluctuate as they do. Answering this seemingly simple question has proven to be challenging. Broadly, density-dependent feedback mechanisms should allow populations to grow at low density and slow or halt growth at high density. However, experimental tests of what demographic processes result in density-dependent feedback and on what timescale have proven elusive. Here, we used replicated density perturbation experiments and capture-mark-recapture analyses to test density-dependent population growth in populations of meadow voles (Microtus pennsylvanicus) during the summer breeding season by manipulating founding population density and observing the pattern of survival, reproduction, and population growth. High population density had no consistent effect on survival rates but generally negatively influenced recruitment and population growth rates. However, these density-dependent effects varied within the breeding season and across years. Our study provides evidence that density-dependent feedback mechanisms operate at finer time scales than previously believed and that process, additively with delayed year effects, is key to understanding multiyear population demography.

Demographic consequences of changes in environmental periodicity.

The fate of natural populations is mediated by complex interactions among vital rates, which can vary within and among years. Although the effects of random, among-year variation in vital rates have been studied extensively, relatively little is known about how periodic, nonrandom variation in vital rates affects populations. This knowledge gap is potentially alarming as global environmental change is projected to alter common periodic variations, such as seasonality. We investigated the effects of changes in vital-rate periodicity on populations of three species representing different forms of adaptation to periodic environments: the yellow-bellied marmot (Marmota flaviventer), adapted to strong seasonality in snowfall; the meerkat (Suricata suricatta), adapted to inter-annual stochasticity as well as seasonal patterns in rainfall; and the dewy pine (Drosophyllum lusitanicum), adapted to fire regimes and periodic post-fire habitat succession. To assess how changes in periodicity affect population growth, we parameterized periodic matrix population models and projected population dynamics under different scenarios of perturbations in the strength of vital-rate periodicity. We assessed the effects of such perturbations on various metrics describing population dynamics, including the stochastic growth rate, log λS . Overall, perturbing the strength of periodicity had strong effects on population dynamics in all three study species. For the marmots, log λS decreased with increased seasonal differences in adult survival. For the meerkats, density dependence buffered the effects of perturbations of periodicity on log λS . Finally, dewy pines were negatively affected by changes in natural post-fire succession under stochastic or periodic fire regimes with fires occurring every 30 years, but were buffered by density dependence from such changes under presumed more frequent fires or large-scale disturbances. We show that changes in the strength of vital-rate periodicity can have diverse but strong effects on population dynamics across different life histories. Populations buffered from inter-annual vital-rate variation can be affected substantially by changes in environmentally driven vital-rate periodic patterns; however, the effects of such changes can be masked in analyses focusing on inter-annual variation. As most ecosystems are affected by periodic variations in the environment such as seasonality, assessing their contributions to population viability for future global-change research is crucial.

Reproductive dispersion and damping time scale with life-history speed.

Iteroparous species may reproduce at many different ages, resulting in a reproductive dispersion that affects the damping of population perturbations, and varies among life histories. Since generation time ( T c ) is known to capture aspects of life-history variation, such as life-history speed, does T c also determine reproductive dispersion ( S ) or damping time ( τ )? Using phylogenetically corrected analyses on 633 species of animals and plants, we find, firstly, that reproductive dispersion S scales isometrically with T c . Secondly, and unexpectedly, we find that the damping time ( τ ) does not scale isometrically with generation time, but instead changes only as T c b with b < 1 (also, there is a similar scaling with S ). This non-isometric scaling implies a novel demographic contrast: increasing generation times correspond to a proportional increase in reproductive dispersion, but only to a slower increase in the damping time. Thus, damping times are partly decoupled from the slow-fast continuum, and are determined by factors other than allometric constraints.

Cooperation by necessity: condition- and density-dependent reproductive tactics of female house mice.

Optimal reproductive strategies evolve from the interplay between an individual's intrinsic state and extrinsic environment, both factors that are rarely fixed over its lifetime. Conditional breeding tactics might be one evolutionary trajectory allowing individuals to maximize fitness. We apply multi-state capture-mark-recapture analysis to a detailed 8-year data set of free-ranging house mice in a growing population to discern causes and fitness consequences of two alternative reproductive tactics in females, communal and solitary breeding. This allows us to integrate natural variation in life-history traits when analysing the expression of two alternative reproductive tactics in females. We find that communal breeding reduces average population fitness, but nevertheless increases over our 8-year study period. The tactic proves to be expressed conditionally dependent on both population density and female body mass - allowing females to breed under subpar conditions, i.e. at high density or when of low body mass. Our results contradict previous laboratory studies and emphasize the importance of studying cooperation under natural conditions, including natural variation in state-dependent survival and breeding probabilities.

Multiple coping strategies maintain stability of a small mammal population in a resource-restricted environment.

In semi-arid environments, aperiodic rainfall pulses determine plant production and resource availability for higher trophic levels, creating strong bottom-up regulation. The influence of climatic factors on population vital rates often shapes the dynamics of small mammal populations in such resource-restricted environments. Using a 21-year biannual capture-recapture dataset (1993 to 2014), we examined the impacts of climatic factors on the population dynamics of the brush mouse (Peromyscus boylii) in semi-arid oak woodland of coastal-central California. We applied Pradel's temporal symmetry model to estimate capture probability (p), apparent survival (φ), recruitment (f), and realized population growth rate (λ) of the brush mouse and examined the effects of temperature, rainfall, and El Niño on these demographic parameters. The population was stable during the study period with a monthly realized population growth rate of 0.993 ± SE 0.032, but growth varied over time from 0.680 ± 0.054 to 1.450 ± 0.083. Monthly survival estimates averaged 0.789 ± 0.005 and monthly recruitment estimates averaged 0.175 ± 0.038. Survival probability and realized population growth rate were positively correlated with rainfall and negatively correlated with temperature. In contrast, recruitment was negatively correlated with rainfall and positively correlated with temperature. Brush mice maintained their population through multiple coping strategies, with high recruitment during warmer and drier periods and higher survival during cooler and wetter conditions. Although climatic change in coastal-central California will likely favor recruitment over survival, varying strategies may serve as a mechanism by which brush mice maintain resilience in the face of climate change. Our results indicate that rainfall and temperature are both important drivers of brush mouse population dynamics and will play a significant role in predicting the future viability of brush mice under a changing climate.

Eco-oncology: Applying ecological principles to understand and manage cancer.

Cancer is a disease of single cells that expresses itself at the population level. The striking similarities between initiation and growth of tumors and dynamics of biological populations, and between metastasis and ecological invasion and community dynamics suggest that oncology can benefit from an ecological perspective to improve our understanding of cancer biology. Tumors can be viewed as complex, adaptive, and evolving systems as they are spatially and temporally heterogeneous, continually interacting with each other and with the microenvironment and evolving to increase the fitness of the cancer cells. We argue that an eco-evolutionary perspective is essential to understand cancer biology better. Furthermore, we suggest that ecologically informed therapeutic approaches that combine standard of care treatments with strategies aimed at decreasing the evolutionary potential and fitness of neoplastic cells, such as disrupting cell-to-cell communication and cooperation, and preventing successful colonization of distant organs by migrating cancer cells, may be effective in managing cancer as a chronic condition.

Contrasting effects of climate change on seasonal survival of a hibernating mammal.

Seasonal environmental conditions shape the behavior and life history of virtually all organisms. Climate change is modifying these seasonal environmental conditions, which threatens to disrupt population dynamics. It is conceivable that climatic changes may be beneficial in one season but result in detrimental conditions in another because life-history strategies vary between these time periods. We analyzed the temporal trends in seasonal survival of yellow-bellied marmots (Marmota flaviventer) and explored the environmental drivers using a 40-y dataset from the Colorado Rocky Mountains (USA). Trends in survival revealed divergent seasonal patterns, which were similar across age-classes. Marmot survival declined during winter but generally increased during summer. Interestingly, different environmental factors appeared to drive survival trends across age-classes. Winter survival was largely driven by conditions during the preceding summer and the effect of continued climate change was likely to be mainly negative, whereas the likely outcome of continued climate change on summer survival was generally positive. This study illustrates that seasonal demographic responses need disentangling to accurately forecast the impacts of climate change on animal population dynamics.

Demography of snowshoe hare population cycles.

Cyclic fluctuations in abundance exhibited by some mammalian populations in northern habitats ("population cycles") are key processes in the functioning of many boreal and tundra ecosystems. Understanding population cycles, essentially demographic processes, necessitates discerning the demographic mechanisms that underlie numerical changes. Using mark-recapture data spanning five population cycles (1977-2017), we examined demographic mechanisms underlying the 9-10-yr cycles exhibited by snowshoe hares (Lepus americanus Erxleben) in southwestern Yukon, Canada. Snowshoe hare populations always decreased during winter and increased during summer; the balance between winter declines and summer increases characterized the four, multiyear cyclic phases: increase, peak, decline, and low. Little or no recruitment occurred during winter, but summer recruitment varied markedly across the four phases with the highest and lowest recruitment observed during the increase and decline phase, respectively. Population crashes during the decline were triggered by a substantial decline in winter survival and by a lack of subsequent summer recruitment. In contrast, initiation of the increase phase was triggered by a twofold increase in summer recruitment abetted secondarily by improvements in subsequent winter survival. We show that differences in peak density across cycles are explained by differences in overall population growth rate, amount of time available for population growth to occur, and starting population density. Demographic mechanisms underlying snowshoe hare population cycles were consistent across cycles in our study site but we do not yet know if similar demographic processes underlie population cycles in other northern snowshoe hare populations.

Every dog has its prey: Range-wide assessment of links between diet patterns, livestock depredation and human interactions for an endangered carnivore.

Livestock depredation is the most ubiquitous type of negative interaction between humans and carnivores. We conducted a range-wide assessment linking diet patterns of the endangered dhole Cuon alpinus, with livestock consumption and human-dhole interactions. We first performed a reanalysis of dhole diet data from all published studies (1973-2013) incorporating a recently-developed non-linear correction factor for quantifying prey biomass consumed. We then determined the relative livestock numbers consumed by dholes over time across its range, compared these with earlier estimates, and investigated the relative importance of wild vs. non-wild prey in dhole diet. Using information from >70 studies, we explored links between livestock consumption by dholes, availability of wild versus non-wild prey, sympatric depredation-prone carnivores, and people's perception of dholes as livestock predators. We found that (a) dhole diet profiles varied regionally, (b) dholes consumed fewer livestock compared to estimates generated using other, widely used methods, (c) livestock consumption by dholes was associated with wild and non-wild prey densities, and number of co-predator species, and (d) people's negative perception of dholes was associated with pack sizes, levels of livestock depredation and number of sympatric carnivore species. Global efforts for dhole conservation should involve different strategies based on region-specific realities that account for ecological context as well as human perceptions, which would require well-designed studies of dhole social and population dynamics, and human-dhole interactions. We also call for more such range-wide assessments of livestock depredation by wild canids, complemented with direct investigations of human-canid interactions.

Assessing seasonal demographic covariation to understand environmental-change impacts on a hibernating mammal.

Natural populations are exposed to seasonal variation in environmental factors that simultaneously affect several demographic rates (survival, development and reproduction). The resulting covariation in these rates determines population dynamics, but accounting for its numerous biotic and abiotic drivers is a significant challenge. Here, we use a factor-analytic approach to capture partially unobserved drivers of seasonal population dynamics. We use 40 years of individual-based demography from yellow-bellied marmots (Marmota flaviventer) to fit and project population models that account for seasonal demographic covariation using a latent variable. We show that this latent variable, by producing positive covariation among winter demographic rates, depicts a measure of environmental quality. Simultaneously, negative responses of winter survival and reproductive-status change to declining environmental quality result in a higher risk of population quasi-extinction, regardless of summer demography where recruitment takes place. We demonstrate how complex environmental processes can be summarized to understand population persistence in seasonal environments.

A critical appraisal of population viability analysis.

Population viability analysis (PVA) is useful in management of imperiled species. Applications range from research design, threat assessment, and development of management frameworks. Given the importance of PVAs, it is essential that they be rigorous and adhere to widely accepted guidelines; however, the quality of published PVAs is rarely assessed. We evaluated the quality of 160 PVAs of 144 species of birds and mammals published in peer-reviewed journals from 1990 to 2017. We hypothesized that PVA quality would be lower with generic programs than with custom-built programs; be higher for those developed for imperiled species; change over time; and be higher for those published in journals with high impact factors (IFs). Each included study was evaluated based on answers to an evaluation framework containing 32 questions reflecting whether and to what extent the PVA study adhered to published PVA guidelines or contained important PVA components. All measures of PVA quality were generally lower for studies based on generic programs. Conservation status of the species did not affect any measure of PVA quality, but PVAs published in high IF journals were of higher quality. Quality generally declined over time, suggesting the quantitative literacy of PVA practitioners has not increased over time or that PVAs developed by unskilled users are being published in peer-reviewed journals. Only 18.1% of studies were of high quality (score >75%), which is troubling because poor-quality PVAs could misinform conservation decisions. We call for increased scrutiny of PVAs by journal editors and reviewers. Our evaluation framework can be used for this purpose. Because poor-quality PVAs continue to be published, we recommend caution while using PVA results in conservation decision making without thoroughly assessing the PVA quality.

Una Evaluación Crítica del Análisis de Viabilidad Poblacional Resumen El análisis de viabilidad poblacional (AVP) es útil para el manejo de especies en peligro. La gama de aplicaciones incluye el diseño de la investigación, la valoración de amenazas y el desarrollo de marcos de trabajo para el manejo. Ya que los AVP son de suma importancia, es esencial que sean rigorosos y se adhieran a las directrices aceptadas por la mayoría; sin embargo, rara vez se examina. la calidad de los AVP publicados Evaluamos la calidad de 160 AVP para 144 especies de aves y mamíferos publicados en revistas con revisión por pares desde 1990 hasta 2017. Nuestra hipotesis consistió en que la calidad del AVP sería más baja con programas genéricos que con programas hechos a la medida; sería más alta para los programas desarrollados para especies en peligro; la calidad cambiaría con el tiempo; y la calidad sería más alta para los AVP publicados en revistas con un alto factor de impacto (VI). Cada estudio que incluimos fue evaluado con base en las respuestas a un marco de trabajo de evaluación que contenía 32 preguntas, las cuales reflejaban si y cuánto se adherían los AVP a las directrices publicadas para los AVP o si contenía componentes importantes de AVP. Todas las medidas de la calidad de los AVP fueron generalmente más bajas para los estudios basados en programas genéricos. El estado de conservación de las especies no afectó ninguna de las medidas de la calidad de los AVP, pero aquellos publicados en revistas con un VI alto tuvieron una mayor calidad. La calidad, en general, declinó con el tiempo, lo que sugiere que el alfabetismo cuantitativo de quienes practican los AVP no ha incrementado con el tiempo o que se están publicando AVP desarrollados por usuarios con poca práctica en revistas con revisión por pares. Sólo el 18.1% de los estudios fue de calidad alta (puntaje >75%), lo cual es preocupante porque los AVP de baja calidad podrían mal informar las decisiones de conservación. Pedimos un incremento en el escrutinio de los AVP por parte de los editores y revisores. Nuestro marco de trabajo de evaluación puede usarse para este propósito. Ya que todavía se publican AVP con baja calidad, recomendamos que se tomen precauciones cuando se usen los resultados de un AVP en la toma de decisiones de conservación sin evaluar minuciosamente la calidad de dicho estudio.

Insights from distribution dynamics inform strategies to conserve a dhole Cuon alpinus metapopulation in India.

Most large carnivore populations currently occur in heterogeneous landscapes, with source populations embedded in a matrix of human-dominated habitats. Understanding changes in distribution of endangered carnivores is critical for prioritizing and implementing conservation strategies. We examined distribution and dynamics of a dhole Cuon alpinus metapopulation, first in 2007 and subsequently in 2015, based on indirect sign surveys across 37, 000sq. km of India's Western Ghats. Predicted dhole occupancy declined from 0.62 (95% CI: 0.58-0.66) in 2007 to 0.54 (95% CI: 0.50-0.58) in 2015. Occupancy was associated with abundance of primary prey species and anthropogenic disturbance. Local extinction appeared to be influenced by forest cover loss, and offset by protected reserves; colonization was influenced by occupancy in neighbouring sites. Perturbation analysis indicated that occupancy was more sensitive to local extinction within reserves and to colonization in sites abutting reserves. The Western Ghats could serve as a stronghold for the endangered dhole, provided future colonizations are facilitated through habitat consolidation beyond reserve boundaries, and local extinctions are prevented by increasing protection efforts within select reserves. We advocate for wildlife managers to adopt a landscape-based approach and periodic monitoring to ensure persistence of the dhole metapopulation in Western Ghats, and in other critical conservation regions across the species' geographic range.