Population Dynamics and Origin of the Himalayan Vulture in Southeast Asia: Phylogeography and Migration Route Analysis

Population Dynamics and Origin of the Himalayan Vulture in Southeast Asia: Phylogeography and Migration Route Analysis | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 29 July 2025 V1 Latest version Share on Population Dynamics and Origin of the Himalayan Vulture in Southeast Asia: Phylogeography and Migration Route Analysis Authors : Chanatip Ummee 0000-0002-2062-6487 , Ratiwan Sitdhibutr , and Chaiyan Kasorndorkbua [email protected] Authors Info & Affiliations https://doi.org/10.22541/au.175374726.69153604/v1 587 views 199 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Since the Industrial Revolution, human activities have led to the decline of vulture species globally. Indian Gyps vultures have experienced a population decline of over 95% due to the drug diclofenac. The Himalayan Vulture ( Gyps himalayensis Hume, 1869) is currently classified as Near Threatened and remains poorly understood owing to its unique high-altitude nesting habits. Migrating from the Tibetan Plateau, the Himalayas, and the Altai Mountains to Southeast Asia in winter, the species often encounters food shortages as it is an obligate scavenger. In this study, satellite transmitters were attached to two Himalayan Vultures (n = 2) between 2021 and 2022. The results revealed that vultures migrating into Thailand returned to their breeding habitats in Sichuan Province, China, and on the Tibetan Plateau. A key finding was that one individual shifted its non-breeding habitat from Southeast Asia to South Asia in the following year. Analysis of home range data showed that the vulture occupied areas at an altitude of 3,300 metres above sea level, with a home range of 291–952 km². The data were then analysed in conjunction with evidence on distribution, evolution, population structure, migration, population dynamics, and efforts related to rehabilitation and release. The findings reveal a population decline on the Qinghai-Tibetan Plateau and in the Indian subcontinent, primarily due to the impacts of diclofenac use and carrion scarcity. In contrast, based on data collected over a 46-year period (1979–2024), a notable population increase has been observed in Southeast Asia, which may result from shifts in migration routes and non-breeding habitats towards the region, likely driven by habitat unsuitability and/or reduced food availability on the Qinghai-Tibetan Plateau due to changes in livestock management practices. These findings suggest that anthropogenic factors may play a crucial role in shaping vulture migration patterns, population dynamics, and even evolutionary trajectories. Since the Industrial Revolution, human activities have led to the decline of vulture species globally. Indian Gyps vultures have experienced a population decline of over 95% due to the drug diclofenac. The Himalayan Vulture ( Gyps himalayensis Hume, 1869) is currently classified as Near Threatened and remains poorly understood owing to its unique high-altitude nesting habits. Migrating from the Tibetan Plateau, the Himalayas, and the Altai Mountains to Southeast Asia in winter, the species often encounters food shortages as it is an obligate scavenger. In this study, satellite transmitters were attached to two Himalayan Vultures (n = 2) between 2021 and 2022. The results revealed that vultures migrating into Thailand returned to their breeding habitats in Sichuan Province, China, and on the Tibetan Plateau. A key finding was that one individual shifted its non-breeding habitat from Southeast Asia to South Asia in the following year. Analysis of home range data showed that the vulture occupied areas at an altitude of 3,300 metres above sea level, with a home range of 291–952 km². The data were then analysed in conjunction with evidence on distribution, evolution, population structure, migration, population dynamics, and efforts related to rehabilitation and release. The findings reveal a population decline on the Qinghai-Tibetan Plateau and in the Indian subcontinent, primarily due to the impacts of diclofenac use and carrion scarcity. In contrast, based on data collected over a 46-year period (1979–2024), a notable population increase has been observed in Southeast Asia, which may result from shifts in migration routes and non-breeding habitats towards the region, likely driven by habitat unsuitability and/or reduced food availability on the Qinghai-Tibetan Plateau due to changes in livestock management practices. These findings suggest that anthropogenic factors may play a crucial role in shaping vulture migration patterns, population dynamics, and even evolutionary trajectories. Keywords: Conservation, Gyps vulture, Population dynamics, Raptor, Systematics, Urbanization Introduction The Himalayan Vulture ( Gyps himalayensis Hume, 1869) is one of eight species in the Gyps genus, which is classified under the Old World vulture group (Subfamily Accipitrinae, Tribe Gypini). This subfamily exhibits characteristics of convergent evolution with another subfamily, Gypaetinae, Tribe Gypaetini (Seibold and Helbig 1995; Wink 1995; Wink and Sauer-Gürth 2004; Lerner and Mindell 2005; Griffiths et al. 2007; Arshad et al. 2009), with their ancestors diverging over 24 million years ago (Mya) (Mindell et al. 2018) . Previous studies suggest that the Gypini vultures is more closely related to other diurnal raptors within the family Accipitridae, such as the Crested Serpent-eagle ( Spilornis cheela Latham, 1790). Within the Gypini Tribe, the first clade consists of Red-headed Vulture ( Sarcogyps calvus Scopoli, 1786), White-headed Vulture ( Trigonoceps occipitalis Burchell, 1824), Lappet-faced Vulture ( Torgos tracheliotos Forster, 1791), and Cinereous Vulture ( Aegypius monachus Linnaeus, 1766), which show close relationships. In contrast, the second clade shows a sister relationship between the Hooded Vulture ( Necrosyrtes monachus Temminck, 1823) and the Gyps vultures. These two clades diverged during the late Miocene around 8 Mya, and a million years later, the ancestors of the Hooded Vulture and Gyp s vultures separated (Johnson et al. 2006; Arshad et al. 2009; Mindell et al. 2018). Gyps vultures originated in the early Pleistocene, around 2 Mya, and form a monophyletic group. Earlier studies using mitochondrial DNA (mtDNA) such as Cytochrome b (Cyt b), NADH dehydrogenase 2 (ND2), and the Control region (CR) suggested that White-rumped Vulture ( Gyps bengalensis Gmelin, 1788) from Asia diverged first (Johnson et al. 2006; Arshad et al. 2009) . However, a more recent multilocus study proposed that White-backed Vulture ( Gyps africanus Salvadori, 1865) from Africa was the first to diverge (Mindell et al. 2018) . The divergence of Himalayan Vulture around 1 Mya is clearer; it separated after White-rumped and White-backed Vultures. It diverged before two clades that display sister relationships: one clade consists of Rüppell’s Vulture ( Gyps rueppelli Brehm, 1852) and Griffon Vulture ( Gyps fulvus Hablizl, 1783), while the other clade consists of Indian Vulture ( Gyps indicus Scopoli, 1786), Slender-billed Vulture ( Gyps tenuirostris Gray, 1844), and Cape Vulture ( Gyps coprotheres Forster, 1798) (Johnson et al. 2006; Arshad et al. 2009; Mindell et al. 2018). These findings reject the previous hypothesis that Himalayan Vulture is closely related to Griffon Vulture ( G. fulvus ) subspecies G. fulvus fulvescens , with which it shares overlapping distribution in some regions (Johnson et al. 2006) (Fig. 1A). Figure 1. Phylogenetic tree and Molecular Dating of the Himalayan Vulture. (A) shows the evolutionary relationships of Old World Vultures, including the divergence timing and speciation of the Himalayan Vulture (Johnson et al. 2006; Arshad et al. 2009; Mindell et al. 2018). The tree topology is based on A: Johnson et al. (2006), B: Arshad et al. (2009), and C: Mindell et al. (2018). (B) illustrates the divergence timing and population separation of Himalayan Vulture (Ummee et al. 2024). Himalayan Vulture was initially studied for population structure using mtDNA Cytb, and the results showed no evidence of geographically separated population structures (Arshad et al. 2009). However, later research revealed that the genetic distance in the CR (Control Region) of the Himalayan Vulture is 10–12 times greater than that in Cytb (Ummee et al. 2023). This led to the use of the CR region to test hypotheses on its population structure. When the CR and Cytb+CR datasets were analysed, a clear population structure emerged, in contrast to the Cytb data, which did not show any distinct structure. Consequently, it was suggested that Cytb is effective for studying species-level relationships, while the CR is better suited for population-level analyses (Ummee et al. 2024), similar to findings in other raptor species (Seibold and Helbig 1995; Wink 1995; Godoy et al. 2004; Lerner and Mindell 2005; Johnson et al. 2006; Hailer et al. 2007; Arshad et al. 2009; Honnen et al. 2010; Langguth et al. 2013). Geographical barriers for Himalayan Vulture include the Trans-Himalayas, Dzungarian Basin (Junggar Basin), Alashan Plateau, Gobi Desert, and Taklamakan Desert. Combined with the natal philopatry behaviour, where raptors or vultures often return to their birthplace to nest and breed (Robertson 1983; Ferrer 1993; Godoy et al. 2004; Hailer et al. 2007; Poulakakis et al. 2008; Honnen et al. 2010; Langguth et al. 2013; Hirschauer et al. 2016; Kleinhans and Willows-Munro 2019), this allowed the species’ population to be divided into three groups: central, western, and northern, or two groups: central and western. The population of Himalayan Vultures diverged approximately 0.2 Mya, during the mid-Pleistocene, before the last glacial period, which occurred between 0.023–0.018 Mya (Ummee et al. 2024) (Fig. 1B and 2). not-yet-known not-yet-known not-yet-known unknown Figure 2. Hypothesis of population divergence in Himalayan Vulture (Ummee et al. 2023, 2024). The black dashed lines represent geographical barriers, and the white arrows indicate the direction of gene flow between populations, analysed using the QGIS software. Molecular dating reveals that the emergence and population divergence of Gyps vultures, including the speciation and population separation of the Himalayan Vulture, occurred during the Early to Middle Pleistocene (2–0.2 Mya). The Pleistocene period was marked by significant climatic fluctuations, with the cycles of glaciations and interglaciations occurring roughly every 100,000 years. These changes in humidity (Koch and Barnosky 2006; Stuart 2015), along with the rise of modern humans (Lyons et al. 2004; Sandom et al. 2014), influenced the evolutionary patterns of various species across the globe, such as speciation, population divergence, and extinction, notably in Africa and Eurasia (Houston 1974; Wilburg and Jackson 1983; Koch and Barnosky 2006; Kendall et al. 2014), as well as in the Americas (Emslie 1998; Stucchi and Emslie 2005; Grayson 2007; Metcalf et al. 2016). The global decrease in temperature and humidity transformed forest canopies into grasslands, which led to an increase in the population of ungulates, particularly in the family Bovidae, which form the primary diet of Gyps vultures (Houston 1974; Houston 1983; Vrba 1985; Arctander et al. 1999; Hassanin and Douzery 1999; Matthee and Davis 2001; Arshad et al. 2009; Perrig et al. 2019). This expansion likely contributed to the spread of the Himalayan Vulture population. Conversely, periods of increasing temperature and humidity resulted in the contraction of the species’ distribution range and the emergence of geographical barriers, limiting gene flow between populations due to the vulture’s specific ecological requirements for nesting at high, cool altitudes (Ummee et al. 2023, 2024). However, this scenario requires further testing with additional markers, such as microsatellites, mitochondrial genomes, single nucleotide polymorphisms (SNPs), or whole-genome analysis, using samples from across the species’ range in mountainous regions surrounding the Qinghai-Tibetan Plateau. The phylogeography of species in these regions is particularly complex (Alcaide et al. 2014; Favre et al. 2015; Dahal et al. 2020; Päckert et al. 2020). In the present Holocene era, Himalayan Vulture faces new challenges, including habitat loss due to rising global temperatures, food scarcity caused by improved public health systems, toxic drug residues from veterinary drugs, poisons used in hunting and livestock protection, and threats from infrastructure development—all of which are linked to human activities. Despite this, the Himalayan Vulture remains one of the least understood vultures, probably due to its specific ecological requirements for nesting at high altitudes, which makes data collection, particularly on its ecology, difficult. However, Thailand’s geographic location places it as a strategic point along the species’ migratory route to Southeast Asia, making it ideal for gathering data, conducting research, and implementing conservation efforts. Long-term studies over the past 15 years have systematically monitored the annual population changes, migration routes, and rehabilitation efforts of the Himalayan Vulture, providing a framework for further research. This review aims to update the public on the ongoing conservation efforts and to assess the feasibility of future studies, ensuring the most effective preparation for the conservation of this globally near-threatened species. Morphology and distribution Himalayan Vulture has a body length of 42 inches, a tail length of 15 inches, a wingspan of 108 inches, and weighs between 8–12 kg, with females being larger than males due to reversed sexual dimorphism. It is the largest species among the eight vultures of the Gyps genus and the second largest of all 9 Asian vultures, particularly in terms of wingspan. Juvenile Himalayan Vultures have buff streaks on a dark brown body, which fade to a cream-white body with contrasting dark brown wing and tail feathers as they mature (Ferguson-Lees and Christie 2001, 2005; Kasorndorkbua et al. 2008). A similar species is the Griffon Vulture, especially in immature plumages, but the Griffon Vulture has lighter rufous-tinted brown body plumage with buff streaks. Additionally, Himalayan Vulture is larger in body size, wingspan, and wing length (Forsman 2016). There are no sighting documentations of the Griffon Vulture in Southeast Asia, whereas Himalayan Vulture regularly winters in this region (Yong and Kasorndorkbua 2008; Sullivan et al. 2009; Subedi 2018; Kasorndorkbua 2024). Himalayan Vultures breed in China, Bhutan, Nepal, India, Pakistan, Afghanistan, Tajikistan, Uzbekistan, Kyrgyzstan, and Kazakhstan (Ali 1962; Inskipp and Inskipp 1985; Ali and Ripley 1968; Thiollay 1994; Knystautas 1993; Grimmett et al. 1999; Mackinnon and Phillipps 2000; Ferguson-Lees and Christie 2001, 2005; Lu et al. 2009; Sullivan et al. 2009; Subedi 2018; Clark et al. 2020; BirdLife International 2024). However, the exact range remains unclear due to the lack of systematic surveys in many countries, such as India (Wani et al. 2020; Kimsing et al. 2021) and Mongolia (BirdLife International 2024; Ummee et al. 2024) (see Fig. 3). One challenge in surveying this species is its nesting behavior, as it builds nests in caves or on cliffs in remote, high-altitude areas, making it difficult to observe and reach. Himalayan Vulture has a highly specialized ecological niche, nesting at high altitudes, making its distribution the one of most restricted of all Old World vultures including Bearded Vulture (Ferguson-Lees and Christie 2001, 2005; MaMing et al. 2013, 2014). Its breeding range follows a C-shape, spanning the Qinghai-Tibetan Plateau, the Himalayas, the Tian Shan Mountains, and the Altai Mountains. In contrast, Griffon Vultures, Cinereous Vultures, and Bearded Vultures, while overlapping in some areas with the Himalayan Vulture, have much broader ranges, extending from Asia and Europe to Africa. The Himalayan Vulture, however, only nests in specific high-altitude regions of Asia (Ferguson-Lees and Christie 2001, 2005; Sullivan et al. 2009; Subedi 2018; Clark et al. 2020; BirdLife International 2024) (see Fig. 3). Figure 3. Distribution range of Himalayan Vulture, including Breeding area (green dashed line) and non-breeding area (red dashed line) (Ferguson-Lees and Christie 2001, 2005; Sullivan et al. 2009; Subedi 2018; Clark et al. 2020; BirdLife International 2024), using QGIS software. Habitat, home range, and migration Himalayan Vulture is a colony breeder. Two to sixteen nests per colony are located on cliff edges or small caves on steep, non-vegetated cliffs (MaMing et al. 2013, 2014; Awan et al. 2017). Nest site selection is positively correlated with proximity to water sources (Wagley et al. 2020) but negatively correlated with human activity areas (Awan et al. 2017). In the Tian Shan Mountain range, Himalayan Vultures build nests at elevations of 2,500–3,200 meters above sea level (MaMing et al. 2013, 2014). One juvenile Himalayan Vulture, ID KU852, fitted with a 25-gram GPS-GSM tracker after rehabilitation, was released back into the wild and tracked in Zamthang County, Sichuan Province, China (Kasorndorkbua et al. 2021). In this study, we analysed the habitat use of KU852 during the breeding season. The habitat was located at an altitude of 3,300 metres above sea level, with a home range estimated between 291 km² (50% kernel density estimate, KDE) and 952 km² (95% KDE) from April 24, 2021, to October 29, 2021, which corresponds to the species’ breeding season (Fig. 4). In contrast, juvenile Himalayan Vultures in the Qinghai-Tibet Plateau, Inner Mongolia, and Mongolia from June to September have an average home range of up to 61,130 km² (±20,062 SE, 95% CI = 21,809–100,452 km²). From November to April, Himalayan Vultures outside the breeding season in India, Nepal, Bhutan, and Tibet have an average home range of 13,973 km² (±6,507 SE, 95% CI = 1,220–26,727 km²) (Sherub et al. 2017). Figure 4. Migration routes and home range of the Himalayan Vulture rescued and released in Thailand. The black lines depict the migration paths of a juvenile Himalayan Vulture with code KU852, including the first spring and second winter seasons in 2021, respectively (Kasorndorkbua et al. 2021). The blue line represents the post-release spring movement of KU903 in 2022 (This study). Both vultures were released at Doi Pha Hom Pok National Park, Chiangmai Province, Thailand along Thailand-Myanmar birder. The white dashed lines indicate periods of missing location data. In this study, we analysed the home range using satellite transmitter tracking data from animal KU852 during the breeding season. The green area represents the home range of KU852 at the 95% kernel density estimate, while the red area represents the home range at the 50% kernel density estimate, as visualised in QGIS. Juvenile Himalayan Vultures undertake long-distance migratory flights from their breeding grounds using two migration routes: the Central Asian Flyway (CAF) to wintering areas in the Indian subcontinent and Central Asia, and the East Asian-Australasian Flyway (EAAF) to Southeast Asia (Kumar et al. 2020) (Fig. 4). According to population separation hypotheses, a larger proportion of Himalayan Vultures migrating to Southeast Asia originates from the Altai Mountains (northern population) and Tian Shan Mountains (western population) (migrating via CAF to EAAF) compared to those from the Qinghai-Tibet Plateau (central population) (migrating via EAAF) (Ummee et al. 2024) (Fig. 2 and 4). Sightings and rescue data in Thailand indicate that Himalayan Vultures use two migration routes into mainland Southeast Asia: the primary route with a higher number of birds goes west, crossing the Thailand-Myanmar border to the Malay Peninsula and Sumatra, Indonesia (Yong and Kasorndorkbua 2008; Bird Society of Singapore 2025). The secondary route, with fewer birds, goes east, entering northern or northeastern Thailand, moving through central and eastern regions, and probably wintering in Cambodia (Kasorndorkbua et al. 2019). There are no reports of sea crossings (Yong and Kasorndorkbua 2008; Sullivan et al. 2009; Kasorndorkbua 2024). However, phylogeographical analysis and satellite tracking have shown that the species’ migration routes to Southeast Asia do not necessarily correlate with the origin of the vultures (Ummee et al. 2024). Non-breeding grounds of Himalayan Vulture include India (Ali 1962; Ali and Ripley 1968; Grimmett et al. 1999; Sullivan et al. 2009; Jha 2015; Subedi 2018; Jha et al. 2020; Praphul and Kumara 2023; BirdLife International 2024), Uzbekistan (Sullivan et al. 2009; Martin et al. 2018), Kazakhstan (Sullivan et al. 2009; Subedi 2018), Turkmenistan (Sullivan et al. 2009), Afghanistan (Sullivan et al. 2009), Iran (Sullivan et al. 2009), Pakistan (Sullivan et al. 2009; Subedi 2018; BirdLife International 2024), UAE (Sullivan et al. 2009), China (Mackinnon and Phillipps 2000; Sullivan et al. 2009; Subedi 2018), Bhutan (Sullivan et al. 2009; Subedi 2018; BirdLife International 2024), Bangladesh (Sullivan et al. 2009; Subedi 2018; BirdLife International 2024), Nepal (Sullivan et al. 2009; Subedi 2018; BirdLife International 2024), Myanmar (Emes 2006; Tordoff et al. 2007; Yong and Kasorndorkbua 2008; Sullivan et al. 2009; Hla et al. 2010; Subedi 2018), Thailand (Round 2007; Kasorndorkbua 2008; Yong and Kasorndorkbua 2008; Sullivan et al. 2009; Kidsin et al. 2012; Subedi 2018; BirdLife International 2024), Cambodia (Gilbert et al. 2006a; Yong and Kasorndorkbua 2008; Sullivan et al. 2009; Subedi 2018; BirdLife International 2024), Malaysia (Jeyarajasingham and Pearson 1999; Wells 1999; Yong and Kasorndorkbua 2008; Sullivan et al. 2009; Chye 2012; Subedi 2018), Singapore (Lim 1998; Wells 1999; Wang and Hails 2007; Yong and Kasorndorkbua 2008; Sullivan et al. 2009; Subedi 2018), and Indonesia (Yong and Kasorndorkbua 2008; Supriatna 2012; Subedi 2018) (Fig. 3). A juvenile Himalayan Vulture with code KU852, fitted with a GPS-GSM tracker and released at Doi Phahom Pok National Park, Chiang Mai, Thailand on April 6, 2021, used the EAAF for its first northward spring migration. It took 24 days to reach its presumed breeding grounds in Sichuan Province, China, where it stayed for over 6 months. On October 29, 2021, KU852 began its second winter migration but switched to the CAF route, eventually passing Tibet, Bhutan, India, and Nepal, arriving in Nepal on November 21, 2021 (Kasorndorkbua et al. 2021) (Table S1 and Fig. 4). Conversely, another juvenile from this study, KU903, also equipped with a 25-gram GSM tracker and released from the same location on April 24, 2022, used the EAAF and was tracked in the Tibetan Plateau from June 15–24, 2022, with missing location data during migration (Table S2 and Fig. 4). Data from satellite tracking of Himalayan Vultures in Thailand, Bhutan, and Nepal reveal that 1) vultures from Sichuan and Tibet migrate between the CAF and EAAF routes, indicating that there are wintering populations in both the Indian subcontinent (India, Nepal, and Bhutan) and Southeast Asia (Sherub et al. 2017; Kasorndorkbua et al. 2021; Subedi 2022), 2) even within the same vulture, there is variation in migration routes and non-breeding habitats as they age (Kasorndorkbua et al. 2021), and 3) there is migration avoiding the high peaks of the Himalayas, also known as the circum-Himalayas route (Sherub et al. 2017; Kasorndorkbua et al. 2021; Subedi 2022). Diet, threats, and breeding success Gyps species primarily consume livestock rather than wildlife carrion. Studies in the Indian subcontinent have shown that western, and northern India consists of livestock (Ghosh-Harihar et al. 2024). Similarly, in the Tibetan Plateau, the diet of Himalayan Vultures is predominantly livestock, such as free-ranging Yak, which makes up 64% of their diet. Wild prey, including wild Yak ( Bos mutus Przewalski, 1883), Tibetan Wild Ass ( Equus kiang Moorcroft, 1841), and Tibetan Antelope ( Pantholops hodgsonii Abel, 1826), constitutes only 1%, and human remains in the ritual of sky burials make up 2% (Lu et al. 2009). However, the proportion of prey may vary according to local human cultural practices. For example, in southern India, Gyps vultures consume up to 77% wild prey, which correlates with higher consumption of cattle and buffalo in that region compared to central, northern, and western India (Ghosh-Harihar et al. 2024). The evolutionary relationship between vultures and humans extends back to the late Pliocene (3.6–2.6 Mya), when early humans, initially scavengers competing with vultures, eventually became hunters who benefited vultures (Moleón et al. 2014). Since the mid-1990s, improvements in human public health have led to a reduction in carrion available to vultures. Concurrently, vultures in the Indian subcontinent faced severe declines due to the use of the Non-steroidal Anti-inflammatory drug (NSAID) Diclofenac in livestock, resulting in a population decline of more than 95–99% (Gilbert et al. 2002; Pain et al. 2003; Oaks et al. 2004; Green et al. 2004; Gilbert et al. 2006b; Swan et al. 2006a, 2006b). Evidence suggests similar impacts on Himalayan Vultures from this drug during their winter migration (Das et al. 2011). In Southeast Asia, the threats to Gyps vultures differ from those in the Indian subcontinent. Major threats include secondary poisoning from pesticides used in hunting or controlling livestock predators, such as Strychnine, Organophosphates, and Carbamates, particularly Carbofuran. These chemicals have led to population declines or local extinctions in certain areas (Clements et al. 2012; Loveridge et al. 2018), such as the extinction of the Red-headed Vulture in Thailand. Food scarcity is another critical factor contributing to the decline of Himalayan Vulture populations. In Azad Jammu and Kashmir, Pakistan, the decline in livestock has significantly affected the Himalayan Vulture population (Siddique and Khan 2016). Similarly, food shortages during winter migration into Southeast Asia, particularly in Thailand due to improved livestock carcass management through slaughterhouses, also contributes to food shortages for the wintering Himalayan Vultures. These have led to consistent mortality due to starvation each year (Kasorndorkbua et al. 2019). There are also reports of vulture deaths from collisions with power lines and poles in Nepal (Subedi 2022). Additionally, climate change may pose a significant threat to Himalayan Vultures, as they inhabit highland areas where rising temperatures are impacting biodiversity (Dahal et al. 2021). Currently, the breeding success rate of Himalayan Vultures ranges between 27–41% from the total number of occupied nests (Karmacharya 2011; Wagley et al. 2020) and 75–90% from active nests (Karmacharya 2011; Joshi et al. 2015). Due to various threats, the conservation status of the Himalayan Vulture has been elevated to Near Threatened since 2014 (BirdLife International 2024). Population dynamics and rehabilitation When Himalayan Vultures are affected by food shortages, they are rescued and sent for veterinary care and rehabilitation at facilities established in various countries. In the Indian subcontinent, there are Wildlife Rescue Centres and Zoos in India (Praveen et al. 2014; Bharos et al. 2020), the Conservation Breeding Centre in National Parks in Nepal (Khadha 2016), and Vulture Rescue Centres in Bangladesh (Alam et al. 2018). In Southeast Asia, Thailand has established a raptor rehabilitation and release program at Kasetsart University Raptor Rehabilitation Unit (KURU), Kasetsart University Veterinary Teaching Hospital Kamphaengsaen with the cooperation of Department of National Parks, Wildlife and Plant Conservation, which provide veterinary cares for rescued or injured Himalayan Vultures and other raptor species (Kasorndorkbua 2024). Since 2005, KURU has rehabilitated 44 Himalayan Vultures and released 36 vultures (all of which were in juvenile age-class when released) back to the wild which a few were tracked for post-release movement from 19 provinces across the country. Similar rescue for release efforts of one Cinereous Vulture, another wintering vulture species have occurred in Singapore as well (Qing 2022). Additionally, there has been a disease monitoring scheme related to bird migration, leading to the first detection of avian malaria, in one juvenile Himalayan Vultures in Thailand (Subaneg et al. 2024). This information is crucial for ongoing monitoring and preparedness for health management and infectious disease prevention in the future. Furthermore, supplementary feeding stations also known as “vulture restaurants” have been established in two provinces, Nakhon Nayok in east-central Thailand and Phuket in southern Thailand, to mitigate food shortages during winter migration (Kasorndorkbua et al. 2019; Kasorndorkbua 2024). These initiatives involve collaborations of birdwatchers, the public, and local authorities, ensuring that carrions used in these restaurants is Diclofenac-free. The species has been benefited from the supplementary feeding strategy that has also been established for the resident species of vultures in Myanmar and Cambodia (Myanmar Vulture Working Group 2019; Cambodia Vulture Working Group 2024). The first documented successful breeding of Himalayan Vultures outside their native range occurred at the Paris Ménagerie in France (Schlee 1989) and later at the Assam State Zoo in India (Ranade et al. 2023). However, there are no captive breeding programs in Southeast Asia. The Himalayan Vulture population is estimated to be between 66,000 and 334,000 individuals (BirdLife International 2024), with up to 229,339±40,447 individuals in the Tibetan Plateau (Lu et al. 2009). Data from sightings and rescues collected from 1979 to 2024 indicate a total of 531 Himalayan Vultures in Southeast Asia. The primary wintering areas are Thailand (327 individuals) and Myanmar (122 individuals), with smaller numbers in Malaysia (16 individuals), Cambodia (7 individuals), and Indonesia (1 individual) (Yong and Kasorndorkbua 2008; Sullivan et al. 2009; Kasorndorkbua 2024) (Fig. 5). This discrepancy may also reflect different coverage in terms of birdwatching activities and surveys in the reported countries. Figure 5. (A) Population Dynamics of Himalayan Vultures Wintering in Southeast Asia from 1979–2024 (Luan-Keng and Hails 2007; Yong and Kasorndorkbua 2008; Sullivan et al. 2009; Kasorndorkbua 2024). Tibet represents population estimates from surveys conducted at Drigung Thel Monastery, Tibet, China, during June to July (Lu et al. 2009). Nepal represents population estimates from surveys conducted in Upper Mustang, Nepal, during July to September (Sharma 2006; Acharya et al. 2009; Paudel et al. 2016). Pakistan represents population estimates from surveys conducted in Azad Jammu and Kashmir, Pakistan, during the breeding season (Siddique and Khan 2016). The dashed line indicates the approximate onset of the diclofenac crisis affecting vultures around 1990, continuing until its ban in India, Nepal, and Pakistan in 2006. Asterisks indicate surveys conducted in breeding habitats, while plus signs indicate surveys conducted in non-breeding habitats. (B) The number of Himalayan Vultures migrating westward through the Thoolakharka raptor migration watch site, Nepal, in 2019, 2020, and 2024 (Gurung et al. 2019; Gurung et al. 2020; Subedi et al. 2025), displayed using R 4.4.1 (R Core Team 2021). The impact of diclofenac from the 1990s onward led to a significant decline in Himalayan Vulture populations across the Indian subcontinent. However, after its ban in 2006, population declines began to stabilize (Gilbert et al. 2002; Galligan et al. 2014). During this period, the population of Himalayan Vultures in their native range of Upper Mustang, Nepal, declined, as observed in surveys conducted before the diclofenac ban in the region (Sharma 2006; Acharya et al. 2009; Paudel et al. 2016). At the same time, food shortages were reported in Azad Jammu and Kashmir, Pakistan, due to a decline in free-range livestock grazing, which led to an increase in shrub cover. This change reduced the availability of carrion and made foraging more difficult, contributing to a decline in the Himalayan Vulture population in the area (Siddique and Khan 2016). Similarly, a decrease in the number of Himalayan Vultures was recorded at Drigung Thel Monastery, Tibet, between 2009 and 2012 (Lu et al. 2009) (Fig. 5). Himalayan Vultures tend to remain in non-breeding areas close to their nesting sites, as evidenced by their almost year-round presence, including during the non-breeding season, in Azad Jammu and Kashmir, Pakistan, which also serves as a breeding ground (Siddique and Khan 2016). In Nepal, they exhibit altitudinal migration, moving from breeding habitats on mountain slopes to wintering grounds in the Terai plains at the foothills of the Himalayas (Rana et al. 2020). Additionally, some breeding habitats also serve as non-breeding areas, as reported on the Qinghai-Tibetan Plateau (Lu et al. 2009; Sherub et al. 2017; Subedi 2022) and in Nepal (Sharma 2006; Virani et al. 2008; Acharya et al. 2009; Paudel et al. 2016; Sherub et al. 2017; Subedi 2022). However, some individuals, particularly juveniles, undertake long-distance movements from breeding grounds (Yong and Kasorndorkbua 2008; BirdLife International 2024). Both the Qinghai-Tibetan Plateau and the Indian subcontinent serve as important non-breeding sites for the species, as confirmed by satellite tracking studies (Sherub et al. 2017; Subedi 2022). Geographical changes caused by human activities, which pose potential threats, may alter the migration routes of vultures. Similar patterns have been observed in raptors such as the Turkey Vulture Cathartes aura (Linnaeus, 1758), Swainson’s Hawk Buteo swainsoni Bonaparte, 1838, and Broad-winged Hawk Buteo platypterus (Vieillot, 1823), which have adjusted their migration routes to avoid increasing numbers of wind farms that pose a danger to them (Cabrera-Cruz and Villegas-Patraca 2016). Likewise, human warfare has also been reported to influence migration patterns of raptors in Europe (Russell et al. 2024). Himalayan Vultures can alter their migration routes as they age. Satellite tracking studies have documented a case of a Himalayan Vulture, likely originating from Sichuan, China that was rescued from starvation in Thailand. This individual shifted its non-breeding site from Southeast Asia i.e., Thailand to South Asia, specifically Nepal, as it aged in the following winter migration (Fig. 4). This change may have resulted from its experience with food scarcity in Southeast Asia, leading to a learned adjustment in migration routes. The impact of diclofenac and food scarcity between the 1990s and 2006 is likely a key factor contributing to the significant increase in the number of Himalayan Vultures in Southeast Asia. The period during which Himalayan Vulture populations declined in South Asia and Tibet (Sharma 2006; Acharya et al. 2009; Lu et al. 2009; Paudel et al. 2016; Siddique and Khan 2016) coincided with a notable rise in their numbers in Southeast Asia, particularly from 2004 onward. The recorded maximum count increased from 9 individuals per year between 1979 and 1992 to as many as 75 individuals per year between 2004 and 2024 (Luan-Keng and Hails 2007; Yong and Kasorndorkbua 2008; Sullivan et al. 2009; Kasorndorkbua 2024) (Fig. 5A). This increase was particularly evident in Myanmar and Thailand, which are located closer to breeding sites and the Eastern Circum-Himalayan Corridor migration route than other countries in the region (Juhant and Bildstein 2017; Ummee et al. 2024). Since 2004, there has been a decline in livestock populations on the Qinghai-Tibet Plateau due to the implementation of Global Warming Potential (GWP) policies, which have led to a reduction in free-range livestock farming (Liu et al. 2021). This may have contributed to the scarcity of carrion for Himalayan Vultures in the region, resulting in an increasing number of individuals migrating to Southeast Asia for the winter since 2004. Surveys at the Thoolakharka raptor migration watch site in Nepal in 2019, 2020, and 2024 recorded an increase in the east-to-west migration of Himalayan Vultures (Gurung et al. 2019; Gurung et al. 2020; Subedi et al. 2025) along the East-to-West Southern Corridor (Juhant and Bildstein 2017) (Fig. 5B). These vultures are likely individuals originating from the Central Population on the Qinghai-Tibetan Plateau (Fig. 2), migrating to non-breeding areas in northwestern India, Pakistan, or Central Asia for the winter. This pattern aligns with the observed decline in breeding-season population estimates on the Qinghai-Tibetan Plateau (Lu et al. 2009) (Fig. 5A). Following the diclofenac crisis, the situation in South Asia has shown signs of improvement, enhancing the region’s capacity to support Himalayan Vultures during the non-breeding season. However, the decline in carrion availability on the Qinghai-Tibetan Plateau remains a growing concern, driven by rising temperatures and humidity, along with a significant reduction in livestock populations from 2001 to 2019 (Liu et al. 2021). This may be linked to the increased migration of vultures from the Qinghai-Tibetan Plateau to northwestern India, Pakistan, or Central Asia between 2019 and 2024 (Gurung et al. 2019; Gurung et al. 2020; Subedi et al. 2025). Following the decline in diclofenac-related impacts, Himalayan Vultures continue to face food shortages in certain habitats, such as the Qinghai-Tibetan Plateau. This has contributed to stable or increasing counts at Thoolakharka, Nepal. This migration route serves as the primary corridor for the central population of Himalayan Vultures, which originates from the Qinghai-Tibetan Plateau and migrates to northwestern India, Pakistan, and Central Asia. In contrast, the population in Southeast Asia, which shows a slight annual increase, follows a secondary migration route with fewer migrating individuals. The shift in migration routes from the Central Asian Flyway (CAF) to the East Asian-Australasian Flyway (EAAF), along with the avoidance of the East-to-West Southern Corridor and the Eastern Circum-Himalayan Corridor due to the impacts of diclofenac and food scarcity, may have facilitated the movement of Himalayan Vultures from their breeding grounds in the Himalayas, including Nepal and Pakistan, as well as from Tibet, Qinghai, and Sichuan in China, along with populations from the Tianshan and Altai Mountains, into Southeast Asia (Fig. 3). It is plausible that these vultures shifted their non-breeding range due to unsuitable conditions in both the Qinghai-Tibet Plateau and the Indian subcontinent. The effects of diclofenac and food scarcity led to continuous population declines at their breeding sites, making these areas less suitable as non-breeding habitats. Consequently, vultures may have undertaken longer migrations to Southeast Asia. Most of these migrants were likely inexperienced, immature individuals that struggled to find food in their native habitats under increasingly harsh conditions, prompting them to migrate farther in search of more favorable foraging opportunities. The impacts of diclofenac and food scarcity are human-induced issues that affect vultures, and they may be key factors driving changes in the migration patterns of Himalayan Vultures between their habitats across the Qinghai-Tibet Plateau, South Asia, and Southeast Asia. This shift in population dynamics could influence the evolutionary patterns that may affect the conservation of this Near Threatened species in the future, especially if regions fail to establish international conservation preparedness, such as the suspension of toxic chemical use, vulture rehabilitation, and reintroduction efforts. Moreover, the creation of supplementary feeding stations to support the vultures’ food requirements especially in winter months could help mitigate food scarcity issues. However, the population increase in Southeast Asia may also be attributed to other contributing factors, such as more systematic data collection and reporting improvements in the region (Yong and Kasorndorkbua 2008; Kasorndorkbua 2024) and more public reporting through broadening citizen science efforts on readily accessible public sighting platforms that enhance reporting efficiency (Sullivan et al. 2009). Additionally, a vulture rehabilitation and release program (Kasorndorkbua et al. 2019), supplementary feeding stations in Thailand, Myanmar, and Cambodia (Hla et al. 2010; Kapetanakos et al. 2014; Kasorndorkbua et al. 2019; Myanmar Vulture Working Group 2019; Cambodia Vulture Working Group 2024). align with the post-release satellite tracking studies (Kasorndorkbua et al. 2021), and phylogeographic research using molecular techniques with samples from Thailand, which currently has the largest dataset (Ummee et al. 2023, 2024), all of which correspond with the increasing trend in Himalayan Vulture numbers in the region. Conclusions and research actions The ecological and evolutionary enigmas surrounding the Himalayan Vulture include its origin in high-altitude regions, geographically structured population, causes of population decline within its native range in contrast to population increases in non-breeding habitats, mysterious migratory routes, population management, and health restoration. These aspects have prompted this study as both a presentation of new findings and a synthesis of the ecological and evolutionary understanding of the Himalayan Vulture. The aim is to provide a comprehensive overview of current research, reveal knowledge gaps, and lay the foundation for future studies. Such studies should promote international collaboration to achieve the most effective conservation outcomes. Research conducted so far and planned for the future will cover the following topics: 1. The distribution range remains uncertain. The Himalayan Vulture’s high-altitude habitat is a key factor contributing to the continued uncertainty surrounding its distribution range. For instance, the northwestern Himalayas in India are included within its global distribution range (BirdLife International 2024). However, no systematic surveys have been conducted in this politically sensitive region. Similarly, parts of the Altai Mountains have been designated as part of the species’ range (BirdLife International 2024), yet there is still no evidence of nesting or breeding activity there, and no comprehensive surveys have been carried out across the area. Phylogeographic studies of the species, when considered alongside migratory routes into Southeast Asia, support the existence of a population in the Altai Mountains (Ummee et al. 2024). This is also backed by satellite telemetry data showing that some Himalayan Vultures migrate to Mongolia during the breeding season, possibly reflecting natal philopatry (Sherub et al. 2017). Therefore, systematic field surveys to confirm the true distribution range of the species are urgently needed. 2. Migration. Data from previous satellite telemetry studies have revealed that the Central Asian Flyway (CAF) and East Asian-Australasian Flyway (EAAF) do not clearly separate Himalayan Vultures migrating from their native range. Additionally, age-related shifts in non-breeding habitats that cross migratory flyways have been observed (Kasorndorkbua et al. 2021). However, satellite tracking data from this study and earlier research in Thailand suggest that individuals wintering in Thailand likely originate from the Himalayas. Still, there is a lack of evidence supporting return migration to the Altai Mountains for vultures using the EAAF to reach Southeast Asia. Therefore, further satellite telemetry tracking is needed, particularly in Thailand, where vultures undergoing rehabilitation due to food shortages are released back into the wild. This approach could clarify whether the Southeast Asian wintering population includes individuals from both the Himalayas and the Altai, supporting the hypothesis of geographically structured populations (Ummee et al. 2024). Such evidence would be critical for informing international conservation policy, especially regarding the management of food scarcity along migratory routes. The increasing number of vultures wintering in Southeast Asia each year makes addressing this issue increasingly urgent. 3. Phylogeography. The hypothesis of geographically structured populations separating the Himalayan, Tien Shan, and Altai mountain ranges has been proposed based on analyses of Himalayan Vultures that migrate to non-breeding areas in Thailand. This hypothesis is supported by genetic evidence in combination with satellite telemetry data from individuals recorded in Thailand during the non-breeding season (Ummee et al. 2023, 2024). These findings highlight an urgent need for international collaboration to collect samples that comprehensively cover all three mountain ranges where nesting is known or suspected to occur. To test this hypothesis with greater resolution, an expanded range of genetic markers should be used, including COI, ND2, Cytb, CR, microsatellites, mitochondrial genomes (mt-genome), SNPs, structural variants (SVs), and whole-genome sequences. This broader genetic approach will enable researchers to uncover the true evolutionary patterns of this elusive and increasingly threatened high-mountain species, providing a robust scientific foundation for its conservation. 4. Threat Identification. Evidence has shown that diclofenac has adverse effects on the Himalayan Vulture (Das et al. 2011). However, since this species breeds on high mountain ranges, the impact of the drug is lower there compared to its wintering grounds in South Asia, where veterinary drug residues are more prevalent. The continuing population decline (BirdLife International 2024), along with the decreasing number of vultures observed near their breeding grounds during the non-breeding season since around 2004 (Sharma 2006; Acharya et al. 2009; Lu et al. 2009; Paudel et al. 2016; Siddique and Khan 2016), contrasts with a marked increase in the number of long-distance migrants arriving in Southeast Asia during the same period. This shift may be linked to the reduction of free-ranging livestock practices across the Tibetan Plateau (Liu et al. 2021), which has led to a decrease in available carrion and may have forced vultures to migrate farther in search of food. Nonetheless, food resources in Southeast Asia remain insufficient. Every year, starving vultures are found collapsed on the ground and require rehabilitation (Kasorndorkbua et al. 2019). This situation underlines the urgent need for further research to confirm food scarcity at both breeding and non-breeding sites. Additionally, international cooperation is required to establish a network of feeding stations along the migration routes, ensuring that carcasses are free from toxic residues. Such coordinated efforts are essential for conserving this species throughout its range. 5. Rehabilitation. In addition to serving as vital facilities for the recovery and subsequent release of undernourished Himalayan Vultures encountered during migration, rehabilitation centres also provide invaluable opportunities for scientific research. These centres serve as important repositories of biological samples such as blood, feathers, and tissue, which are essential for evolutionary studies (Ummee et al. 2023, 2024). They also support ecological research through satellite tagging to study migratory patterns (Kasorndorkbua et al. 2021), as well as veterinary studies and clinical interventions based on direct treatment and handling of individual birds (Kidsin et al. 2012; Subaneg et al. 2024; Sitdhibutr et al. 2025). Such research would be extremely difficult to conduct in the high-altitude breeding areas where these vultures originate. For this reason, the establishment of rehabilitation centres in non-breeding grounds provides conservation benefits in both the short term, by restoring the health of individuals for release into the wild, and in the long term, by enabling the collection of biological materials for ecological, evolutionary, and veterinary studies. It is therefore an urgent priority to support the development of rehabilitation units in all countries where cases of starving vultures have been reported along the migratory route. This will foster international cooperation and enhance the overall effectiveness of conservation efforts for this threatened highland species. In conclusion, food scarcity and the use of diclofenac have contributed to a decline in the populations of Himalayan Vulture on the Qinghai-Tibet Plateau and in South Asia, the number of these vultures migrating to Southeast Asia for the non-breeding season increased significantly. As a result, this region may have become important wintering grounds, compensating for the loss of suitable or less food-available habitats in the Qinghai-Tibet Plateau and South Asia. With over a decade-long of data collection and ongoing efforts in monitoring and rehabilitation for release programs of the species, it is evident that mainland Southeast Asia plays a crucial role in the conservation of this species along the EAAF. 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Keywords gyps vulture conservation population dynamics raptor systematics urbanization Authors Affiliations Chanatip Ummee 0000-0002-2062-6487 Kasetsart University Faculty of Veterinary Medicine View all articles by this author Ratiwan Sitdhibutr Kasetsart University Veterinary Teaching Hospital Kamphaeng Saen View all articles by this author Chaiyan Kasorndorkbua [email protected] Kasetsart University Faculty of Veterinary Medicine View all articles by this author Metrics & Citations Metrics Article Usage 587 views 199 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Chanatip Ummee, Ratiwan Sitdhibutr, Chaiyan Kasorndorkbua. Population Dynamics and Origin of the Himalayan Vulture in Southeast Asia: Phylogeography and Migration Route Analysis. Authorea . 29 July 2025. DOI: https://doi.org/10.22541/au.175374726.69153604/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. 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