Megaherbivores provide biotic resistance against alien plant dominance.

While human-driven biological invasions are rapidly spreading, finding scalable and effective control methods poses an unresolved challenge. Here, we assess whether megaherbivores-herbivores reaching ≥1,000 kg of body mass-offer a nature-based solution to plant invasions. Invasive plants are generally adapted to maximize vegetative growth. Megaherbivores, with broad dietary tolerances, could remove large biomass of established plants, facilitating new plant growth. We used a massive dataset obtained from 26,838 camera stations and 158,979 vegetation plots to assess the relationships between megaherbivores, native plants and alien plants across India (~121,330 km2). We found a positive relationship between megaherbivore abundance and native plant richness and abundance, and a concomitant reduction in alien plant abundance. This relationship was strongest in protected areas with midproductive ecosystem and high megaherbivore density but it was lost in areas where thicket-forming alien plants predominated (>40% cover). By incorporating the role of ecosystem productivity, plants traits and densities of megaherbivores on megaherbivore-vegetation relationships, our study highlights a function of megaherbivores in controlling alien plant proliferation and facilitating diverse native plants in invaded ecosystems. The study shows great potential for megafauna-based trophic rewilding as a nature-based solution to counteract dominance of plant invasions.

Synthesizing habitat connectivity analyses of a globally important human-dominated tiger-conservation landscape.

As ecological data and associated analyses become more widely available, synthesizing results for effective communication with stakeholders is essential. In the case of wildlife corridors, managers in human-dominated landscapes need to identify both the locations of corridors and multiple stakeholders for effective oversight. We synthesized 5 independent studies of tiger (Panthera tigris) connectivity in central India, a global priority landscape for tiger conservation, to quantify agreement on landscape permeability for tiger movement and potential movement pathways. We used the latter analysis to identify connectivity areas on which studies agreed and stakeholders associated with these areas to determine relevant participants in corridor management. Three or more of the 5 studies' resistance layers agreed in 63% of the study area. Areas in which all studies agree on resistance were of primarily low (66%, e.g., forest) and high (24%, e.g., urban) resistance. Agreement was lower in intermediate resistance areas (e.g., agriculture). Despite these differences, the studies largely agreed on areas with high levels of potential movement: >40% of high average (top 20%) current-flow pixels were also in the top 20% of current-flow agreement pixels (measured by low variation), indicating consensus connectivity areas (CCAs) as conservation priorities. Roughly 70% of the CCAs fell within village administrative boundaries, and 100% overlapped forest department management boundaries, suggesting that people live and use forests within these priority areas. Over 16% of total CCAs' area was within 1 km of linear infrastructure (437 road, 170 railway, 179 transmission line, and 339 canal crossings; 105 mines within 1 km of CCAs). In 2019, 78% of forest land diversions for infrastructure and mining in Madhya Pradesh (which comprises most of the study region) took place in districts with CCAs. Acute competition for land in this landscape with globally important wildlife corridors calls for an effective comanagement strategy involving local communities, forest departments, Appendix 1 and infrastructure planners. This article is protected by copyright. All rights reserved.

Camera trap placement for evaluating species richness, abundance, and activity.

Information from camera traps is used for inferences on species presence, richness, abundance, demography, and activity. Camera trap placement design is likely to influence these parameter estimates. Herein we simultaneously generate and compare estimates obtained from camera traps (a) placed to optimize large carnivore captures and (b) random placement, to infer accuracy and biases for parameter estimates. Both setups recorded 25 species when same number of trail and random cameras (n = 31) were compared. However, species accumulation rate was faster with trail cameras. Relative abundance indices (RAI) from random cameras surrogated abundance estimated from capture-mark-recapture and distance sampling, while RAI were biased higher for carnivores from trail cameras. Group size of wild-ungulates obtained from both camera setups were comparable. Random cameras detected nocturnal activities of wild ungulates in contrast to mostly diurnal activities observed from trail cameras. Our results show that trail and random camera setup give similar estimates of species richness and group size, but differ for estimates of relative abundance and activity patterns. Therefore, inferences made from each of these camera trap designs on the above parameters need to be viewed within this context.

Contrasting levels of social distancing between the sexes in lions.

Understanding sexual segregation is crucial to comprehend sociality. A comparative analysis of long-term lion data from Serengeti and Ngorongoro in Tanzania, and Gir in India, reveals that male-female associations are contingent upon male and female group size, prey-size and availability, and the number of prides that each male coalition currently resides. Males maintain proximity with females, whereas females are responsible for segregation except at large kills. Lions feed on the largest prey in Ngorongoro and the smallest in Gir, and females spend the most time with males in Ngorongoro and the least in Gir. Females roar less often in prey-scarce circumstances in Serengeti and throughout the year in Gir possibly to prevent being tracked by males that parasitize on female kills. However, females readily associate with males when available prey is large and abundant. Contrasting availability of resources between Gir and Serengeti/Ngorongoro helps explain the varying degrees of sexual segregation and appears to drive differences in mating systems between these lion populations.

Feasibility of reintroducing grassland megaherbivores, the greater one-horned rhinoceros, and swamp buffalo within their historic global range.

Reintroduction of endangered species is an effective and increasingly important conservation strategy once threats have been addressed. The greater one-horned rhinoceros and swamp buffalo have declined through historic hunting and habitat loss. We identify and evaluate available habitat across their historic range (India, Nepal, and Bhutan) for reintroducing viable populations. We used Species Distribution Models in Maxent to identify potential habitats and evaluated model-identified sites through field visits, interviews of wildlife managers, literature, and population-habitat viability analysis. We prioritize sites based on size, quality, protection, management effectiveness, biotic pressures, and potential of conflict with communities. Our results suggest that populations greater than 50 for rhinoceros and 100 for buffalo were less susceptible to extinction, and could withstand some poaching, especially if supplemented or managed as a metapopulation. We note some reluctance by managers to reintroduce rhinoceros due to high costs associated with subsequent protection. Our analysis subsequently prioritised Corbett and Valmiki, for rhino reintroduction and transboundary complexes of Chitwan-Parsa-Valmiki and Dudhwa-Pilibhit-Shuklaphanta-Bardia for buffalo reintroductions. Establishing new safety-nets and supplementing existing populations of these megaherbivores would ensure their continued survival and harness their beneficial effect on ecosystems and conspecifics like pygmy hog, hispid hare, swamp deer, hog deer, and Bengal florican.

Recent Evolutionary History of Tigers Highlights Contrasting Roles of Genetic Drift and Selection.

Species conservation can be improved by knowledge of evolutionary and genetic history. Tigers are among the most charismatic of endangered species and garner significant conservation attention. However, their evolutionary history and genomic variation remain poorly known, especially for Indian tigers. With 70% of the world's wild tigers living in India, such knowledge is critical. We re-sequenced 65 individual tiger genomes representing most extant subspecies with a specific focus on tigers from India. As suggested by earlier studies, we found strong genetic differentiation between the putative tiger subspecies. Despite high total genomic diversity in India, individual tigers host longer runs of homozygosity, potentially suggesting recent inbreeding or founding events, possibly due to small and fragmented protected areas. We suggest the impacts of ongoing connectivity loss on inbreeding and persistence of Indian tigers be closely monitored. Surprisingly, demographic models suggest recent divergence (within the last 20,000 years) between subspecies and strong population bottlenecks. Amur tiger genomes revealed the strongest signals of selection related to metabolic adaptation to cold, whereas Sumatran tigers show evidence of weak selection for genes involved in body size regulation. We recommend detailed investigation of local adaptation in Amur and Sumatran tigers prior to initiating genetic rescue.

The role of kinship and demography in shaping cooperation amongst male lions.

The influence of kinship on animal cooperation is often unclear. Cooperating Asiatic lion coalitions are linearly hierarchical; male partners appropriate resources disproportionately. To investigate how kinship affect coalitionary dynamics, we combined microsatellite based genetic inferences with long-term genealogical records to measure relatedness between coalition partners of free-ranging lions in Gir, India. Large coalitions had higher likelihood of having sibling partners, while pairs were primarily unrelated. Fitness computations incorporating genetic relatedness revealed that low-ranking males in large coalitions were typically related to the dominant males and had fitness indices higher than single males, contrary to the previous understanding of this system based on indices derived from behavioural metrics alone. This demonstrates the indirect benefits to (related) males in large coalitions. Dominant males were found to 'lose less' if they lost mating opportunities to related partners versus unrelated males. From observations on territorial conflicts we show that while unrelated males cooperate, kin-selected benefits are ultimately essential for the maintenance of large coalitions. Although large coalitions maximised fitness as a group, demographic parameters limited their prevalence by restricting kin availability. Such demographic and behavioural constraints condition two-male coalitions to be the most attainable compromise for Gir lions.

GPS-telemetry unveils the regular high-elevation crossing of the Himalayas by a migratory raptor: implications for definition of a "Central Asian Flyway".

Remote technologies are producing leapfrog advances in identifying the routes and connectivity of migratory species, which are still unknown for hundreds of taxa, especially Asian ones. Here, we used GPS-telemetry to uncover the migration routes and breeding areas of the massive population of migratory Black-eared kites wintering around the megacity of Delhi-India, which hosts the largest raptor concentration of the world. Kites migrated for 3300-4800 km along a narrow corridor, crossing the Himalayas at extremely high elevations (up to > 6500 m a.s.l.) by the K2 of the Karakoram Range and travelled long periods at elevations above 3500 m. They then crossed/circumvented the Taklamakan Desert and Tian Shan Range to reach their unknown breeding quarters at the intersection between Kazakhstan, Russia, China and Mongolia. Route configuration seemed to be shaped by dominant wind support and barrier avoidance. Wintering ranges were smaller than breeding ranges and concentrated around Delhi, likely in response to massive human food-subsidies. Our results illustrate that high-elevation crossings by soaring migrants may be more common than previously appreciated and suggest the delineation of a hitherto poorly-appreciated "Central Asian Flyway", which must funnel hundreds of thousands of migrants from central Asia into the Indian subcontinent via multiple modes of the Himalayan crossing.

Spatially explicit density and its determinants for Asiatic lions in the Gir forests.

Asiatic lions (Panthera leo persica) are an icon of conservation success, yet their status is inferred from total counts that cannot account for detection bias and double counts. With an effort of 4,797 km in 725 km2 of western Gir Protected Area, India, we used polygon search based spatially explicit capture recapture framework to estimate lion density. Using vibrissae patterns and permanent body marks we identified 67 lions from 368 lion sightings. We conducted distance sampling on 35 transects with an effort of 101.5 km to estimate spatial prey density using generalized additive modeling (GAM). Subsequently, we modeled lion spatial density with prey, habitat characteristics, anthropogenic factors and distance to baiting sites. Lion density (>1-year-old lions) was estimated at 8.53 (SE 1.05) /100 km2 with lionesses having smaller movement parameter (σ = 2.55 km; SE 0.12) compared to males (σ = 5.32 km; SE 0.33). Detection corrected sex ratio (female:male lions) was 1.14 (SE 0.02). Chital (Axis axis) was the most abundant ungulate with a density of 63.29 (SE 10.14) as determined by conventional distance sampling (CDS) and 58.17 (SE 22.17)/km2 with density surface modeling (DSM), followed by sambar (Rusa unicolor) at 3.84 (SE 1.07) and 4.73 (SE 1.48)/km2 estimated by CDS and DSM respectively. Spatial lion density was best explained by proximity to baiting sites and flat valley habitat but not as much by prey density. We demonstrate a scientifically robust approach to estimate lion abundance, that due to its spatial context, can be useful for management of habitat and human-lion interface. We recommend this method for lion population assessment across their range. High lion densities in western Gir were correlated with baiting. The management practice of attracting lions for tourism can perturb natural lion densities, disrupt behavior, lion social dynamics and have detrimental effects on local prey densities.

The population density of an urban raptor is inextricably tied to human cultural practices.

Human socio-cultural factors are recognized as fundamental drivers of urban ecological processes, but their effect on wildlife is still poorly known. In particular, human cultural aspects may differ substantially between the extensively studied urban settings of temperate regions and the poorly studied cities of the tropics, which may offer profoundly different niches for urban wildlife. Here, we report how the population levels of a scavenging raptor which breeds in the megacity of Delhi, the black kite Milvus migrans, depend on spatial variation in human subsidies, mainly in the form of philanthropic offerings of meat given for religious purposes. This tight connection with human culture, which generated the largest raptor concentration in the world, was modulated further by breeding-site availability. The latter constrained the level of resource-tracking by the kites and their potential ecosystem service, and could be used as a density-management tool. Similar ties between animal population densities, key anthropogenic resources and human beliefs may occur in thousands of cities all over the globe and may fit poorly with our current understanding of urban ecosystem functioning. For many urban animals, key resources are inextricably linked with human culture, an aspect that has been largely overlooked.

Human-attacks by an urban raptor are tied to human subsidies and religious practices.

Growing urbanization is increasing human-wildlife interactions, including attacks towards humans by vertebrate predators, an aspect that has received extremely scarce investigation. Here, we examined the ecological, landscape and human factors that may promote human-aggression by raptorial Black kites Milvus migrans in the 16-millions inhabitants megacity of Delhi (India). Physical attacks depended on human activities such as unhygienic waste management, ritual-feeding of kites (mainly operated by Muslims), human density, and presence of a balcony near the nest, suggesting an association between aggression and frequent-close exposure to humans and derived food-rewards. Surprisingly, while more than 100,000 people could be at risk of attack in any given moment, attitudes by local inhabitants were strikingly sympathetic towards the birds, even by injured persons, likely as a result of religious empathy. These results highlight the importance of socio-cultural factors for urban biota and how these may radically differentiate the under-studied cities of developing countries from those of western nations, thus broadening our picture of human-wildlife interactions in urban environments. The rapid sprawling of urban and suburban areas with their associated food-subsidies is likely to increase proximity and exposure of large predators to humans, and vice versa, leading to heightened worldwide conflicts.

Offspring defense by an urban raptor responds to human subsidies and ritual animal-feeding practices.

There is a growing interest in the behavioural and life history mechanisms that allow animal species to cope with rapidly expanding urban habitats, which impose frequent proximity to humans. A particular case of behavioral bottleneck (i.e. conflicting interests) faced by animals in urban environments is how they will modulate the defence of their offspring against the potential danger represented by humans, an aspect that has received scarce research attention. We examined the nest defense against humans by a dense breeding population of a raptor, the Black Kite Milvus migrans, within the megacity of Delhi (India). Here, kites live on a diet dominated by human waste and meat offered through religiously motivated bird feeding practices. Nest defense levels increased with the number of offspring, and with the progression of the breeding season. Defense also intensified close to ritual-feeding areas and with increasing human waste in the streets, suggesting synergistic effects of food availability, parental investment, personality-boldness and habituation to humans, with consequent attenuation of fear. Thus, the behavioural response to a perceived threat reflected the spatial mosaic of activity of humans in the city streets, their cultural practices of ritual-feeding, and their waste-management. For synurbic species, at the higher-end spectrum of adaptation to an urban life, human cultural practices and attitudes may well be the most defining dimensions of their urban niche. Our results suggest that, after initial urban colonization, animals may continue to adapt to the typically complex, heterogeneous environments of cities through fine-grained behavioural adjustments to human practices and activities.

Seasonal dynamics in mosquito abundance and temperature do not influence avian malaria prevalence in the Himalayan foothills.

We examined seasonal prevalence in avian haemosporidians (Plasmodium and Haemoproteus) in migrant and resident birds in western Himalaya, India. We investigated how infection with haemosporidians in avian hosts is associated with temporal changes in temperature and mosquito abundance along with host abundance and life-history traits (body mass). Using molecular methods for parasite detection and sequencing partial cytochrome b gene, 12 Plasmodium and 27 Haemoproteus lineages were isolated. Our 1-year study from December 2008 to December 2009 in tropical Himalayan foothills revealed a lack of seasonal variation in Plasmodium spp. prevalence in birds despite a strong correlation between mosquito abundance and temperature. The probability of infection with Plasmodium decreased with increase in temperature. Total parasite prevalence and specifically Plasmodium prevalence showed an increase with average avian body mass. In addition, total prevalence exhibited a U-shaped relationship with avian host abundance. There was no difference in prevalence of Plasmodium spp. or Haemoproteus spp. across altitudes; parasite prevalence in high-altitude locations was mainly driven by the seasonal migrants. One Haemoproteus lineage showed cross-species infections between migrant and resident birds. This is the first molecular study in the tropical Himalayan bird community that emphasizes the importance of studying seasonal variation in parasite prevalence. Our study provides a basis for further evolutionary study on the epidemiology of avian malaria and spread of disease across Himalayan bird communities, which may not have been exposed to vectors and parasites throughout the year, with consequential implications to the risk of infection to naïve resident birds in high altitude.

Conservation priorities for endangered Indian tigers through a genomic lens.

Tigers have lost 93% of their historical range worldwide. India plays a vital role in the conservation of tigers since nearly 60% of all wild tigers are currently found here. However, as protected areas are small (<300 km2 on average), with only a few individuals in each, many of them may not be independently viable. It is thus important to identify and conserve genetically connected populations, as well as to maintain connectivity within them. We collected samples from wild tigers (Panthera tigris tigris) across India and used genome-wide SNPs to infer genetic connectivity. We genotyped 10,184 SNPs from 38 individuals across 17 protected areas and identified three genetically distinct clusters (corresponding to northwest, southern and central India). The northwest cluster was isolated with low variation and high relatedness. The geographically large central cluster included tigers from central, northeastern and northern India, and had the highest variation. Most genetic diversity (62%) was shared among clusters, while unique variation was highest in the central cluster (8.5%) and lowest in the northwestern one (2%). We did not detect signatures of differential selection or local adaptation. We highlight that the northwest population requires conservation attention to ensure persistence of these tigers.

Selfish partners: resource partitioning in male coalitions of Asiatic lions.

Behavioral plasticity within species is adaptive which directs survival traits to take multiple pathways under varying conditions. Male-male cooperation is an evolutionary strategy often exhibiting an array of alternatives between and within species. African male lions coalesce to safeguard territories and mate acquisition. Unique to these coalitions is lack of strict hierarchies between partners, who have similar resource securities possibly because of many mating opportunities within large female groups. Skewed mating and feeding rights have only been documented in large coalitions where males were related. However, smaller modal prey coupled with less simultaneous mating opportunities for male Asiatic lions in Gir forests, India would likely result in a different coalition structure. Observations on mating events (n = 127) and feeding incidents (n = 44) were made on 11 male coalitions and 9 female prides in Gir, to assess resource distribution within and among different sized male coalitions. Information from 39 males was used to estimate annual tenure-holding probabilities. Single males had smaller tenures and appropriated fewer matings than coalition males. Pronounced dominance hierarchies were observed within coalitions, with one partner getting more than 70% of all matings and 47% more food. Competition between coalition partners at kills increased with decline in prey size, increase in coalition size and the appetite states of the males. However, immediate subordinates in coalitions had higher reproductive fitness than single males. Declining benefits to partners with increasing coalition size, with individuals below the immediate subordinates having fitness comparable to single males, suggest to an optimal coalition size of 2 lions. Lions under different competitive selection in Gir show behavioral plasticity to form hierarchical coalitions, wherein partners utilize resources asymmetrically, yet coalesce for personal gains.

Human Perceptions Mirror Realities of Carnivore Attack Risk for Livestock: Implications for Mitigating Human-Carnivore Conflict.

Human-carnivore conflict is challenging to quantify because it is shaped by both the realities and people's perceptions of carnivore threats. Whether perceptions align with realities can have implications for conflict mitigation: misalignments can lead to heightened and indiscriminant persecution of carnivores whereas alignments can offer deeper insights into human-carnivore interactions. We applied a landscape-scale spatial analysis of livestock killed by tigers and leopards in India to model and map observed attack risk, and surveyed owners of livestock killed by tigers and leopards for their rankings of threats across habitats to map perceived attack risk. Observed tiger risk to livestock was greatest near dense forests and at moderate distances from human activity while leopard risk was greatest near open vegetation. People accurately perceived spatial differences between tiger and leopard hunting patterns, expected greater threat in areas with high values of observed risk for both carnivores. Owners' perception of threats largely did not depend on environmental conditions surrounding their village (spatial location, dominant land-use or observed carnivore risk). Surveys revealed that owners who previously lost livestock to carnivores used more livestock protection methods than those who had no prior losses, and that owners who had recently lost livestock for the first time expressed greater interest in changing their protection methods than those who experienced prior losses. Our findings suggest that in systems where realities and perceptions of carnivore risk align, conservation programs and policies can optimize conservation outcomes by (1) improving the effectiveness of livestock protection methods and (2) working with owners who have recently lost livestock and are most willing to invest effort in adapting protection strategies to mitigate human-carnivore conflict.

Ancient Himalayan wolf (Canis lupus chanco) lineage in Upper Mustang of the Annapurna Conservation Area, Nepal.

The taxonomic status of the wolf (Canis lupus) in Nepal's Trans-Himalaya is poorly understood. Recent genetic studies have revealed the existence of three lineages of wolves in the Indian sub-continent. Of these, the Himalayan wolf, Canis lupus chanco, has been reported to be the most ancient lineage historically distributed within the Nepal Himalaya. These wolves residing in the Trans-Himalayan region have been suggested to be smaller and very different from the European wolf. During October 2011, six fecal samples suspected to have originated from wolves were collected from Upper Mustang in the Annapurna Conservation Area of Nepal. DNA extraction and amplification of the mitochondrial (mt) control region (CR) locus yielded sequences from five out of six samples. One sample matched domestic dog sequences in GenBank, while the remaining four samples were aligned within the monophyletic and ancient Himalayan wolf clade. These four sequences which matched each other, were new and represented a novel Himalayan wolf haplotype. This result confirms that the endangered ancient Himalayan wolf is extant in Nepal. Detailed genomic study covering Nepal's entire Himalayan landscape is recommended in order to understand their distribution, taxonomy and, genetic relatedness with other wolves potentially sharing the same landscape.