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Boselaphus

Boselaphus is a genus of bovid in the family Bovidae, subfamily Bovinae, and tribe Boselaphini, comprising the largest antelope native to Asia as its only extant species, the nilgai (Boselaphus tragocamelus), along with at least one extinct species, B. namadicus, known from the Pliocene and Pleistocene deposits of the Siwalik Group in South Asia.^[1] The genus, first established by Blainville in 1816 with B. tragocamelus (originally described by Pallas in 1766) as the type species, represents a Miocene relict lineage characterized by primitive horn morphology and adaptation to open, dry habitats.^[1] The nilgai, meaning "blue bull" in Hindi, is a sexually dimorphic ungulate endemic to the Indian subcontinent, where it inhabits grasslands, woodlands, and scrublands from the foothills of the Himalayas southward to the peninsula, including parts of India, Nepal, and northeastern Pakistan.^[2]^[1] Adult males stand 120–150 cm at the shoulder, weigh 180–288 kg, and exhibit a striking blue-gray coat with short, smooth, black horns (15–24 cm long), a distinctive throat bib, and chest shield, while females are smaller (up to 212 kg), tawny brown, and hornless.^[1] These herbivores are diurnal browsers and grazers, consuming grasses, leaves, and fruits, and typically form small groups of fewer than 10 individuals, with males often solitary outside the breeding season; they can reach speeds of 48 km/h when fleeing predators such as tigers or dholes.^[1] Fossil evidence indicates that Boselaphus originated in the late Miocene and diversified in South Asia, with B. namadicus—larger than the modern nilgai—documented from 3.4 to 0.6 million years ago in the Upper Siwaliks, including finds of dentition and postcranial elements from Pleistocene sites in Pakistan's Pabbi Hills.^[1]^[3] The genus is distinguished from related boselaphines like the four-horned antelope (Tetracerus) by its size, lack of anterior horns, and compressed neck morphology.^[1] As of 2008, the nilgai population exceeded 100,000 in India alone and is classified as Least Concern by the IUCN, though it faces local threats from habitat loss and is considered an agricultural pest due to crop raiding; introduced populations thrive in Texas, USA, since the 1920s.^[2]^[1]

Etymology and taxonomy

Name origin

The genus name Boselaphus is derived from the Latin word bos, meaning "ox" or "cow," combined with the Greek elaphos, meaning "deer," highlighting the animal's robust, bovine body structure paired with more slender, cervine traits.^[4]^[5] The species epithet tragocamelus originates from the Greek words tragos ("he-goat") and kamelos ("camel"), a reference to the male's horned profile resembling a goat and its sloping back akin to a camel's.^[5] The nilgai was first scientifically described by German zoologist Peter Simon Pallas in 1766 under the name Antilope tragocamelus, based on accounts from northeastern India.^[1] In 1816, French zoologist Henri Marie Ducrotay de Blainville established the genus Boselaphus and transferred the species to it as the type.^[6] The common name "nilgai" stems from the Hindi words nil ("blue") and gāy ("cow"), alluding to the slate-blue coat of adult males.^[7]

Classification history

The genus Boselaphus was established by French zoologist Henri Marie Ducrotay de Blainville in 1816, with the nilgai (Boselaphus tragocamelus) designated as the type species; this species had previously been described by Peter Simon Pallas in 1766 under the name Antilope tragocamelus.^[8]^[9] Early classifications placed the nilgai under various genera, including Antilope and occasionally Capra, reflecting the limited understanding of bovid diversity at the time.^[9] Synonyms for the genus include Portax (proposed by Andrew Smith in 1827, often applied to fossil forms resembling the nilgai) and, in some historical contexts, subgenera under Tragelaphus as defined by Blainville in 1816.^[10] Within the family Bovidae, Boselaphus is classified in the subfamily Bovinae and the tribe Boselaphini, which distinguishes it from related tribes such as Tragelaphini (spiral-horned antelopes) and Bovini (cattle and allies) based on morphological traits like horn structure and cranial features.^[1] The tribe Boselaphini traces its formal recognition to divisions outlined by John Edward Gray in 1846, who grouped Boselaphus and similar forms (e.g., under Strepsicereae) separate from other bovines, though the tribe name itself was later solidified in classifications like those by George Gaylord Simpson in 1945.^[11]^[12] Taxonomic debates at the subtribe and genus levels have occasionally proposed lumping Boselaphus with Tetracerus (the four-horned antelope) into a broader Boselaphini, given shared primitive bovid characteristics, but Boselaphus has been maintained as distinct to accommodate nilgai-like forms with straight, keeled horns and robust builds.^[1]^[13] Molecular analyses since the early 2000s, incorporating mitochondrial DNA (e.g., cytochrome b) and nuclear markers, have robustly confirmed Boselaphus as a monotypic genus within Bovinae, with the nilgai as its sole extant species and a basal position relative to other bovine tribes.^[14] These studies, including phylogenetic reconstructions from 2003 onward, underscore the genus's evolutionary isolation and support its current taxonomic stability.^[15]

Phylogenetic relationships

Boselaphus is classified within the order Artiodactyla, suborder Ruminantia, family Bovidae, subfamily Bovinae, and tribe Boselaphini.^[16] The tribe Boselaphini forms a sister group to the tribes Bovini (including cattle and buffaloes) and Tragelaphini (including kudu and eland), with their divergence estimated at approximately 10–12 million years ago during the Miocene epoch.^[17]^[18] Genetic analyses utilizing mitochondrial DNA (mtDNA) and nuclear loci, particularly from studies in the 2010s, position Boselaphus as basal relative to more derived bovine tribes, with the four-horned antelope (Tetracerus quadricornis) identified as its closest living relative within Boselaphini.^[16]^[18] Morphological characteristics supporting this phylogenetic placement include keeled horn cores in males, absence of accessory (false) hooves, and specialized features of the ruminant digestive system adapted for mixed feeding, which distinguish Boselaphini from other bovine tribes.^[19]^[20] The lineage of Boselaphus connects to early Miocene boselaphines, such as forms akin to Eotragus, highlighting its retention of primitive traits among extant bovids.^[17]

Species

Boselaphus tragocamelus

Boselaphus tragocamelus, commonly known as the nilgai or blue bull, is the sole extant species in the genus Boselaphus and serves as its type species.^[1] As the largest antelope native to Asia, it inhabits open landscapes across the Indian subcontinent, where it plays a key role in grassland ecosystems as a herbivore. Boselaphus tragocamelus is considered monotypic, with no recognized subspecies. The species exhibits pronounced sexual dimorphism, with males displaying a characteristic bluish-gray coat and females a tawny brown, adaptations that aid in camouflage within their preferred environments.^[5]^[21] Physically, the nilgai reaches a shoulder height of 120-150 cm, with adult males weighing 180-288 kg and females 100-212 kg.^[1] It prefers grasslands and scrublands of the Indian subcontinent, including areas from the Himalayan foothills to southern India, while avoiding dense forests.^[5]^[22] Behaviorally, it is diurnal, functioning as both a grazer and browser, primarily consuming grasses and woody plants, and typically forms small herds of 2-18 individuals, though larger groups occur occasionally.^[5]^[21] When threatened, it can flee at speeds up to 50 km/h.^[5] The nilgai is classified as Least Concern by the IUCN, with a wild population estimated at over 100,000 individuals, primarily in India.^[2] It has been introduced to Texas, USA, since the 1920s, where feral populations now number around 30,000-50,000.^[23]^[24]

Boselaphus namadicus

Boselaphus namadicus is the only known extinct species within the genus Boselaphus, classified as a member of the tribe Boselaphini based on shared cranial and dental features with the modern nilgai. This bovid inhabited South Asia during the Late Pliocene to Early Pleistocene, with a temporal range spanning approximately 3.4 to 0.6 million years ago, primarily documented from the Siwalik Group sediments. Fossils have been recovered from sites in India, including the Siwalik Hills and Narmada Valley, and in Pakistan from the Pabbi Hills and Sardhok formations, indicating a distribution across the Indo-Pakistani subcontinent in diverse fluvial and alluvial environments.^[25] The species was first described in the late 19th century from Indian fossils, initially named Portax namadicus by Rütimeyer in 1878 before being reassigned to Boselaphus by Lydekker in the same year due to morphological affinities with the genus. Early discoveries focused on cranial material from the Narmada basin's older alluvium, representing Middle Pleistocene deposits. More recent excavations, such as those in 2017 from the Sardhok Pleistocene outcrops in northern Pakistan, have yielded additional cranial and postcranial elements, including mandibular fragments with deciduous premolars and molars, as well as isolated teeth, enhancing understanding of its variability. These finds from the Pabbi Hills also include the first reported upper deciduous premolar from the Pleistocene Siwaliks, confirming the species' presence in the region during the early Middle Pleistocene.^[25]^[26] Fossil evidence reveals B. namadicus as a large-sized bovid, larger than the extant B. tragocamelus, with estimated body masses exceeding 300 kg based on postcranial scaling. Cranial morphology includes horn cores positioned closer to the orbits, featuring a more pronounced inner keel directed inward and forward, straighter overall structure compared to modern relatives, wider occipital condyles, and a larger foramen magnum. Postcranial elements suggest robust limbs adapted for open terrains, while dental features comprise hypsodont, narrow-crowned molars and trilobed deciduous premolars with crescentic lobes; measurements from Pakistani fossils include dp4 lengths of 29.0–31.7 mm and m1 widths up to 13.8 mm, indicating a robust dentition suited to abrasive vegetation.^[25] The extinction of B. namadicus occurred towards the end of the Middle Pleistocene, around 0.6 million years ago, likely driven by climate shifts towards drier conditions, habitat loss from faunal turnover in the Siwalik region, and increased competition with other bovids. This event coincided with broader environmental changes in South Asia, marking the disappearance of several large mammals from the Pleistocene assemblages.^[25]

Physical characteristics

Morphology and size

Boselaphus species exhibit a robust body structure typical of large bovids, featuring a sloping back that descends from the shoulders to the hindquarters, a deep neck with a prominent white throat patch, and a short erectile mane or crest running along the nape and back. The legs are slender and thin, adapted for agile movement and speed across open terrains, supporting a stocky frame. The head is long and narrow, with a short, coarse coat that varies from blue-gray in adult males to tawny-brown in females and juveniles.^[5]^[21] Adults of the genus typically measure 1.2–1.5 m at the shoulder, with body lengths ranging from 170–210 cm and tail lengths of 40–45 cm; weights vary from 169–288 kg, with males generally larger and heavier than females at 200–250 kg on average compared to 150–200 kg for females. These dimensions position Boselaphus as Asia's largest antelope genus, emphasizing their adaptation to expansive habitats.^[5]^[21]^[1] Horns are present exclusively in males, forming short, cylindrical structures that are ringed, diverge outward, and curve gently backward; they average 15–24 cm in length, with maximums up to 25 cm, and lack the pronounced rings seen in many other bovids.^[5]^[23]^[1] Sensory adaptations include well-developed eyesight and hearing, comparable to or exceeding that of sympatric deer species, aiding in predator detection over distances; the sense of smell is less acute. Large eyes support effective vision in varying light conditions, though the genus is primarily diurnal with some crepuscular activity.^[5]^[7] Fossil representatives, such as the Pleistocene Boselaphus namadicus, display morphological variations including larger overall body mass and distinct horn-core features, such as more inwardly keeled and forward-directed bases, compared to the standardized form of the extant B. tragocamelus; dental measurements indicate proportionally larger teeth, with lower molars up to 32 mm in length.^[25]^[27]

Sexual dimorphism and adaptations

Boselaphus species exhibit pronounced sexual dimorphism, particularly in B. tragocamelus, where males are larger and possess distinct morphological features adapted for territorial interactions. Adult males develop a blue-gray coat, contrasting with the tawny or yellowish-brown coloration of females, which aids in camouflage within grassy habitats.^[28]^[5] Males also feature a prominent throat beard up to 13 cm long beneath a white gular patch, along with black mane on the forequarters, while females have a rudimentary or absent beard.^[28]^[23] This dimorphism extends to body size, with males averaging 241 kg and reaching up to 288 kg, compared to females at 169 kg on average and up to 212 kg, enabling males to engage in physical defense of territories.^[28] Only males bear horns, which are short, straight, sharp, and black, measuring 15–24 cm in length with occasional basal rings indicating growth increments; these structures are absent or rudimentary in females.^[28]^[5] The horns serve a functional role in male-male combat, supported by a thick dermal shield on the neck and chest that provides protection during confrontations.^[28]^[5] Females, being smaller and more agile, lack these robust features, aligning with evasion-based survival strategies.^[23] Juveniles of both sexes resemble females in appearance, with a pale brown or tawny coat that persists until physical maturity around 2–3 years of age, after which males transition to the adult blue-gray coloration.^[5]^[23] This similarity aids in predator avoidance during early development. Beyond dimorphism, B. tragocamelus displays adaptations for arid environments, including a thin, oily coat and the ability to derive most moisture from vegetation, allowing survival for extended periods without direct water intake.^[5] In the extinct B. namadicus, similar sexual dimorphism is inferred from fossil cranial and horn morphology, where variations in size and structure suggest male-dominant horn development for intraspecific competition, akin to the modern species.^[29]

Distribution and ecology

Native range and habitat preferences

The genus Boselaphus, represented primarily by its extant species B. tragocamelus (nilgai), is native to the Indian subcontinent, encompassing central and northern India, southern Nepal, and parts of Pakistan.^[30] Historically, the range extended into northwestern Bangladesh, though the species was extirpated there by the early 20th century. As of 2024, at least 15 individuals have been sighted in 7 districts since 2018 through incursions from neighboring India, though some have been poached and conservation efforts face challenges, including the death of a rescued female in 2025.^[31]^[32]^[33] The nilgai's distribution favors open landscapes across elevations from sea level to approximately 1,800 meters, particularly in the foothills of the Himalayas extending southward to regions like Mysore in southern India.^[5] Nilgai inhabit a variety of dry to semi-arid environments, including grassy steppes, scrub forests, floodplains, and dry deciduous woodlands, often along agricultural edges where human-modified landscapes provide foraging opportunities.^[34] They prefer open grasslands with scattered short trees and bushes, avoiding dense jungles while tolerating arid conditions; however, they require seasonal access to water sources such as rivers or waterholes, as populations are influenced by water availability and may shift territories during prolonged dry periods.^[35] These habitats are typically in monsoon-influenced tropical climates with temperatures ranging from 20°C to 40°C, supporting the species' adaptations to seasonal variations in vegetation and precipitation.^[36] Microhabitat use involves daytime resting in shaded areas under trees or scrub for thermoregulation, transitioning to open grassy patches for nighttime grazing, which aligns with their preference for undulating terrain that balances cover and visibility.^[23] This pattern underscores the genus's ecological niche in transitional zones between forests and plains, distinct from fully desert or closed-canopy environments.^[37]

Introduced populations

The nilgai (Boselaphus tragocamelus) was first introduced to the United States in 1924, when individuals were imported from zoos in India and released on private ranches in southern Texas around 1930.^[7]^[13] These early translocations established feral populations that have since expanded across semi-arid regions of South Texas, particularly on large ranches in counties such as Kenedy, Kleberg, and Willacy, where the species thrives due to suitable open grasslands and brushy habitats similar to its native range.^[7]^[36] As of 2021, the introduced population in Texas was estimated at approximately 37,000 individuals, supported by the absence of natural predators and ample forage resources.^[38] Populations have also become established in northeastern Mexico through natural dispersal from Texas and direct introductions, with nilgai now present in Tamaulipas and adjacent areas, where they occupy similar arid and semi-arid landscapes.^[36]^[39] Other introduced populations exist in South Africa on large ranches (>85,000 ha) and historically in Italy near Rome, though the latter disappeared during World War II.^[1] In introduced ranges, nilgai exhibit minimal interbreeding with native ungulates due to genetic and behavioral barriers, maintaining their distinct lineage.^[13] Ecologically, introduced nilgai compete with native white-tailed deer (Odocoileus virginianus) for forage in shared rangelands, potentially reducing deer densities in high-overlap areas, though overall impacts on native biodiversity appear limited based on available studies.^[40]^[41] They also serve as vectors for cattle fever ticks (Rhipicephalus spp.), posing risks to livestock health and prompting disease surveillance efforts.^[42]^[43] Nilgai are actively hunted as exotic game on private lands, contributing to population control and economic benefits for ranchers through guided hunts.^[44] In the United States, nilgai receive no legal protection and are classified as non-native exotics, allowing landowners to manage them via hunting or culling without permits, particularly to mitigate disease transmission risks such as bovine tuberculosis and tick-borne pathogens.^[7]^[40] Similar management approaches apply in Mexico, where populations are monitored for livestock health implications but not conserved as wildlife.^[39]

Behavior

Boselaphus species exhibit sexually segregated social structures outside the breeding season, with females and calves forming stable matriarchal herds typically comprising 4–15 individuals, often led by a dominant female that directs group movements and decisions.^[45] These family units provide protection for young and facilitate cooperative vigilance against predators. Mature males, in contrast, are primarily solitary or associate in bachelor groups of 2–18 individuals, where subadult and non-breeding adults engage in low-intensity social interactions to practice dominance behaviors.^[45]^[36] Within these groups, hierarchies are pronounced, particularly among males, who establish dominance through aggressive encounters involving horn clashes, head butting, and neck wrestling, which can escalate to serious injuries or even mortality during intense rivalries.^[45] Female hierarchies are less overt but involve displays such as rushing and butting to maintain order within family units, ensuring efficient resource access and calf safety.^[45] Communication among individuals relies on a combination of vocalizations, including guttural grunts and alarm bleats to signal threats or coordinate group activities, alongside olfactory cues from preorbital gland secretions and conspicuous dung piles used for scent marking and advertisement.^[45]^[36] Territoriality is most evident in mature males during the rut (October–February in native ranges), when solitary individuals defend personal space through repeated visits to latrine sites and aggressive displays toward intruders, while non-breeding periods see more nomadic movements across larger home ranges.^[36]^[7] Outside the breeding season, both sexes exhibit fluid group fission-fusion dynamics influenced by resource availability, though overall gregariousness remains moderate compared to more social bovids.^[36]

Diet and foraging

Boselaphus species are herbivorous, with diets primarily consisting of grasses and browse. In populations of B. tragocamelus in Texas, the diet comprises approximately 60% grasses, 25% forbs, and 15% browse, though this can shift to include more browse during periods of scarcity.^[46] In Indian habitats, the composition is more balanced toward browse, with grasses making up about 42%, fallen leaves, flowers, and fruits at 36%, browse at 17%, and herbs or shrubs at 6%, utilizing over 120 plant species including crops such as maize when available near agricultural areas.^[47]^[48] Foraging in B. tragocamelus is predominantly diurnal, with activity concentrated in morning and late-afternoon peaks that may extend into crepuscular periods, totaling around 8-10 hours per day depending on season and food availability.^[49] During the wet season, individuals engage in selective grazing on fresh grasses, while in the dry season, they shift to browsing on leaves, twigs, and fruits to meet nutritional needs.^[46] Crop raiding occurs seasonally in agricultural zones, particularly targeting maize and other grains, which supplements the natural diet but leads to conflicts with farmers.^[48] Additionally, nilgai seek out mineral licks for salts and essential nutrients, frequenting natural soil licks rich in sodium and other minerals.^[50] The digestive system of Boselaphus features a four-chambered stomach adapted for rumination, enabling efficient breakdown of fibrous plant material through microbial fermentation and regurgitation.^[13] This physiology supports survival in arid conditions, as B. tragocamelus can endure 2-3 days without direct water intake by deriving moisture metabolically from vegetation.^[49] Fossil evidence from B. namadicus indicates a similar mixed-feeding strategy, with dental wear patterns suggesting consumption of both grasses and browse in open, drier Pleistocene habitats.^[37]

Reproduction and life history

Mating systems

Boselaphus tragocamelus exhibits a polygynous mating system in which a single mature male mates with multiple females, with dominance areas rather than fixed harems or territories established during the breeding season.^[1] In native ranges across India and Pakistan, the rut peaks from October to December, though breeding activity extends from October to February overall, with males actively seeking receptive females during this period.^[1] Courtship is relatively simple and brief, often lasting around 45 minutes, involving the male approaching the female with a stiff-legged gait and erect tail while vocalizing with grunts or a peculiar mooing sound to signal interest.^[1] Males also engage in scent marking through preorbital gland rubbing on vegetation and urination to advertise dominance, alongside aggressive displays such as threatening postures, neck wrestling, and chasing rivals to secure access to females.^[21] If receptive, the female raises her tail, allowing the male to lick or nuzzle her perineum before mounting.^[1] Gestation lasts approximately 240 days, resulting in the birth of usually one calf, though twinning occurs in some populations.^[1] During the rut, males show heightened aggression toward competitors and reduced feeding, leading to significant energy expenditure and weight loss.^[21] Sexual dimorphism, with males larger and possessing horns absent in females, is evident in extant Boselaphus and inferred for extinct congeners from Miocene and Pleistocene fossils, indicating similar polygynous systems drove the evolution of these traits in the genus.^[1]^[51] During breeding, temporary groups form around dominant males and several females, facilitating mate access.^[1]

Development and lifespan

Nilgai calves are typically born after a gestation period of 243–247 days, with females usually producing a single offspring, although twins occur in up to 56% of pregnancies in introduced populations in Texas (as of 2024) and are reported as common in some native studies.^[52]^[53] Newborn calves weigh approximately 6–16 kg depending on the source and population, and they are precocial, able to stand and walk within 20–40 minutes of birth.^[54]^[55] Mothers nurse their young from a reverse parallel position, and calves begin showing interest in solid forage by the fourth week of life.^[52] Weaning occurs around 10 months of age, after which young males typically leave their mothers to join bachelor groups.^[21] Sexual maturity is reached by females at approximately 18–24 months in native populations, though few breed before 3 years, while males mature later at 3–4 years and do not effectively compete for mates until around 4–5 years; in introduced populations in Texas, females can reach maturity as early as 1 year old (as of 2024).^[52]^[53] Individuals attain full adult size by 3–4 years, with males continuing to develop their characteristic blue-gray coat and robust build into early adulthood.^[52] In the wild, nilgai have a lifespan of 12–13 years, though high juvenile mortality rates of around 34% annually contribute to lower average longevity, with many calves not surviving their first year.^[52] In captivity, they can live up to 21 years.^[52] Aging is indicated by annual rings on horn bases and tooth wear, allowing age estimation up to about 9–10 years; older individuals show worn horns and may exhibit faded coat coloration, but there is no distinct menopause, as females can reproduce into advanced age.^[52]

Fossil record and evolution

Miocene origins

The tribe Boselaphini originated in the early Miocene, approximately 18–20 million years ago, from early boselaphine stock that had dispersed across Eurasia and possibly Africa.^[56] Ancestral forms include the basal boselaphin Miotragocerus, known from middle to late Miocene deposits in Central Europe and Asia, dating to the transition between the Vallesian and Turolian stages around 12–7 Ma, which exhibits plesiomorphic dental and postcranial features indicative of early boselaphine morphology.^[57] These lineages trace back further to primitive bovids like Eotragus, a small, horned form from the early Miocene around 18 Ma, representing one of the earliest bovid genera with true horn cores.^[58] Migration of early boselaphines to Asia occurred via the Tethys Seaway during the Miocene, facilitating their spread from African origins into Eurasian habitats.^[59] A key divergence within the Bovinae subfamily saw Boselaphini split from the sister tribe Tragelaphini around 12 Ma, with the Boselaphini clade becoming well-established by 7 Ma as evidenced by diverse late Miocene fossils.^[60] This separation is supported by molecular phylogenies placing Boselaphini and Tragelaphini as a close cluster within Bovinae, diverging from other tribes 10–15 Ma ago.^[56] Innovations during this period included the evolution of larger body sizes compared to early Miocene ancestors, enabling adaptation to more open environments, alongside horn core developments such as basal insertion and keeling in forms like Miotragocerus, which differed from the more spiral forms in Tragelaphini.^[61] These evolutionary changes occurred amid the late Miocene climatic cooling, which promoted the expansion of C3-dominated grasslands across Eurasia and Africa, favoring boselaphine adaptations for browsing and grazing in mixed woodland-grassland mosaics.^[62] Stable isotope analyses of late Miocene boselaphine teeth confirm dietary shifts toward C3 vegetation in these emerging open habitats, underscoring their ecological flexibility during this transition.^[63]

Pliocene and Pleistocene fossils

Fossil records of Boselaphus in the Pliocene are primarily known from early forms resembling B. namadicus within the Siwalik Group sediments of India and Pakistan, dating to approximately 3.4–2.6 million years ago (Ma), with the genus likely evolving from late Miocene boselaphines such as Pachyportax in the Siwalik region. These remains consist mainly of isolated teeth and horn cores, such as the fourth upper deciduous premolar (DP4), which provide evidence of the genus's presence in Late Pliocene fluvial environments.^[25] During the Pleistocene, Boselaphus fossils have been documented at several key sites, including the Kurnool Caves in southern India, where remains of Boselaphus sp. indicate Late Pleistocene occupation alongside other ungulates. In Pakistan, excavations at Pabbi Hills, including those conducted around 2017, have yielded bovid material attributed to B. namadicus, while the Sardhok locality has produced more complete skeletal elements, such as mandibles and dentition, dated to approximately 0.6 Ma.^[64]^[25]^[27] These Pleistocene fossils often occur in riverine deposits associated with fauna such as elephants (Elephas namadicus) and hippopotamuses (Hexaprotodon sp.), reflecting a paleoecological setting of woodland-grassland mosaics along ancient fluvial systems in the Siwalik region.^[65] Taphonomic analysis of Boselaphus bones from these sites reveals evidence of predation, including tooth puncture marks, gnaw damage, and spiral fractures, which suggest that herds were vulnerable to large carnivores contributing to fossil accumulation in stream-influenced contexts.^[66] A 2018 discovery of the related boselaphine genus Duboisia from Pleistocene deposits in Thailand indicate a broader historical distribution of the tribe across Asia beyond the Indian subcontinent.^[67]

Conservation

Population status

The global population of the nilgai (Boselaphus tragocamelus), the sole living species in the genus Boselaphus, exceeds 100,000 individuals in India, where numbers are considered stable or increasing, with introduced populations in Texas, USA, estimated at around 50,000.^[2]^[44] These figures reflect a recovery from severe declines in the 19th century due to intensive hunting, bolstered by legal protections under India's Wildlife Protection Act of 1972, which classifies the nilgai as a Schedule III species.^[68] Populations in native ranges outside India, such as Pakistan and Nepal, remain small and fragmented. In prime habitats across India, nilgai densities typically range from 10 to 40 individuals per km², with higher concentrations in protected areas such as Gir Forest where local estimates exceed 40 per km² in some western Indian landscapes.^[69] For example, Gujarat hosts one of the largest subpopulations, with over 250,000 individuals recorded in 2021, representing a 117% increase over the previous decade.^[68] Introduced populations in Texas, USA, number 40,000 to 60,000 and continue to grow at an annual rate of 5-10%, primarily along the southern border regions where they were first established in the 1920s from zoo escapes and private ranches.^[44] Monitoring of nilgai populations in India relies on methods such as camera traps for occupancy modeling and line transect surveys to estimate abundance and density, particularly in tiger reserves and agricultural fringes where human-wildlife interactions are common.^[70] These techniques, often integrated with distance sampling and GIS analysis, provide data for state-level censuses but lack comprehensive national coordination, leading to reliance on aggregated regional estimates. No formal population counts exist for extinct fossil species within Boselaphus, as assessments focus on the extant nilgai.

Threats and protection

The primary threats to Boselaphus tragocamelus, commonly known as the nilgai or blue bull, stem from anthropogenic activities that have significantly reduced its native habitat across the Indian subcontinent. Habitat fragmentation and loss, primarily driven by agricultural expansion and urbanization, have led to a substantial contraction of the species' range due to conversion of grasslands and scrublands into farmlands.^[71]^[34] In regions like Nepal and Pakistan, this fragmentation exacerbates isolation of populations, increasing vulnerability to local extirpations.^[71]^[72] Poaching for meat, hides, and horns remains a persistent risk, particularly in areas bordering protected zones where enforcement is limited, contributing to population declines in unprotected landscapes.^[34]^[73] Human-wildlife conflict further compounds these pressures, as nilgai frequently raid crops in agricultural fields, leading to retaliatory killings and culling by farmers; in states like Bihar and Uttar Pradesh in India, such conflicts have prompted localized declarations of the species as vermin to allow controlled culling. As of 2025, rising populations have intensified these conflicts, prompting further culling approvals in northern Indian states.^[74]^[71]^[75] In native ranges, natural predation by tigers (Panthera tigris) and leopards (Panthera pardus) can impact juveniles and weakened individuals, though this is less significant compared to human-induced threats.^[73] Disease transmission poses additional risks, particularly in introduced populations outside the native range. In the southern United States, where nilgai were introduced in the 1920s, there is potential for disease spillover from domestic cattle, although direct infections in nilgai remain unconfirmed and surveillance focuses on monitoring interspecies contact. Climate change exacerbates these vulnerabilities through increased drought frequency, which reduces forage availability in arid grasslands and may lead to nutritional stress and higher mortality rates during dry seasons.^[71] Conservation efforts for B. tragocamelus include legal protections and habitat management to mitigate these threats. In India, the species is listed under Schedule III of the Wildlife (Protection) Act, 1972, which restricts hunting and trade but allows for control measures in conflict zones.^[76] Globally, it is included in CITES Appendix III (via Pakistan), regulating international trade to prevent overexploitation.^[77] Key protected areas in India, such as Ranthambore National Park, Gir National Park, and Bandipur National Park, support stable populations through anti-poaching patrols and habitat restoration, covering thousands of hectares of suitable grassland ecosystems.^[78]^[79] In introduced ranges like Texas, where nilgai populations have expanded to over 50,000 individuals, management emphasizes regulated hunting to control numbers and prevent overgrazing or disease spread; year-round hunting with no bag limits is permitted under state licenses, generating revenue for habitat conservation while maintaining ecological balance.^[80]^[81] These measures, combined with ongoing monitoring, help sustain the species' overall least concern status on the IUCN Red List, though localized declines underscore the need for continued enforcement against habitat loss and conflict.^[82]

References

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Boselaphus Tragocamelus (Artiodactyla: Bovidae) - Oxford Academic
Boselaphustragocamelus (Pallas, 1766) is a bovid commonly called the nilgai or blue bull and is Asia's largest antelope. A sexually dimorphic ungulate of large ...
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The IUCN Red List of Threatened Species
**Summary of Boselaphus tragocamelus (Nilgai)**
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BOSELAPHUS Definition & Meaning - Merriam-Webster
Boselaphus. noun. Bo·sel·a·phus. bōˈzeləfəs. : a genus of Asian antelopes including only the nilgai. Word History. Etymology. New Latin, from Latin bos ox, ...
[4]
Boselaphus tragocamelus (nilgai) - Animal Diversity Web
Nilgai antelope are among the largest of the asian antelopes. They stand about 120-150cm at the shoulder and have a body length of 180-200cm.
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Mammal Species of the World - Browse: Boselaphus
GENUS Boselaphus. Author: de Blainville, 1816. Citation: Bull. Sci. Soc. Philom. Paris, 1816: 75. Type Species: Antilope tragocamelus Pallas, 1766. EXPORT AS ...
[6]
Nilgai Antelope: History, Habitat, and Management in Texas
Nilgai antelope (Boselaphus tragocamelus Pallas) were apparently brought to the United States from India as zoo animals before the mid-1920s and were ...
[7]
Boselaphus Blainville, 1816 - GBIF
Genus Accepted. Boselaphus Blainville, 1816. Published in: Bull. Sci. Soc. Philom. Paris vol.1816 p.75. In: GBIF Backbone Taxonomy · 2,605 occurrences.
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Mammal Species of the World - Browse: tragocamelus
Antilope tragocamelus Pallas, 1766, was based on accounts of the "tragelaphus" by Caius, Gesner and Ray and on Parsons' (1745) description of a male nilgai ...
[9]
Genus Boselaphus - Nomenclature & Taxonomy - The Taxonomicon
Authorship, types, synonyms, homonyms, common names, taxonomic positions and number of subtaxa of Genus Boselaphus Blainville, 1816.<|separator|>
[10]
Review of Family-Group Names of Living Bovids - Oxford Academic
Until their validity is confirmed, Boselaphini, Tragelaphini, Cephalophinae, and Alcelaphinae also are to be employed in preference to their senior synonyms. If ...
[11]
The Oldest Bovid from the Siwaliks, Pakistan - jstor
Tribe BOSELAPHINI Simpson, 1945. Genus EOTRAGUS Pilgrim, 1939. Type Species -Eotragus sansaniensis (Lart6t, 1851). Generic Diagnosis--The combination of six mo.
[12]
Boselaphus Tragocamelus (Artiodactyla: Bovidae) - BioOne Complete
Jun 6, 2008 · Boselaphus tragocamelus (Pallas, 1766) is a bovid commonly called the nilgai or blue bull and is Asia's largest antelope.
[13]
Phylogenetic reconstruction and the identification of ancient ...
Apr 24, 2009 · We identified 3 distinct lineages after the Bovini split from the Boselaphini and Tragelaphini tribes, which has lead to the (1) Buffalo ...Missing: history | Show results with:history
[14]
Molecular and Morphological Phylogenies of Ruminantia and the ...
Abstract.—The ruminants constitute the largest group of ungulates, with >190 species, and its distribution is widespread throughout all continents except ...
[15]
A multi-calibrated mitochondrial phylogeny of extant Bovidae ...
Phylogenetic studies based on genomic data provide an unparalleled tool for the reconstruction of phylogenetic relationships among organisms. However ...
[16]
Tribal Origins and Radiations
Boselaphini is a large, diverse, and, as currently understood, non-monophyletic group. A revision of the systematics of taxa variously grouped as Boselaphini is ...
[17]
Phylogenetic reconstruction and the identification of ancient ...
Apr 24, 2009 · We identified 3 distinct lineages after the Bovini split from the Boselaphini and Tragelaphini tribes, which has lead to the (1) Buffalo ...
[18]
Total Evidence Phylogenetic Analysis Supports New Morphological ...
Jun 15, 2021 · The ultimate goal of this study was to find hard-tissue synapomorphies for the family Bovidae by optimizing morphological characters on a total ...
[19]
Molecular Insights into the Evolution of the Family Bovidae
The combined nuclear DNA data proved to be informative in recovering evolutionary relationships among some bovid subfamilies (fig. 2 ). The Bovidae are ...
[20]
Nilgai - Ultimate Ungulate
Nilgai is from the Hindustani word nil, meaning blue, and the Persian word gaw, meaning cow: a reference to the male's slate blue colouration. Bos (Latin) an ox ...<|separator|>
[21]
A species account of the Nilgai (Boselaphus tragocamelus) | TTU
Nilgai are large antelopes, native to India and Pakistan, with males having short horns. They are gray to brownish-gray with white markings, and are found in ...
[22]
Boselaphus tragocamelus (Pallas, 1766) - GBIF
Classified as Least Concern on The IUCN Red List. There are about 100,000 Nilgai in India, very few in Pakistan, and they were extirpated from Bangladesh ...
[23]
[PDF] Latrine ecology of nilgai antelope - East Foundation
Nilgai antelope (Boselaphus tragocamelus) are native to India,. Nepal, and Pakistan. They were introduced to Texas rangelands in the United States during the ...
[24]
Fossils of Boselaphus (Bovini: Bovidae: Ruminantia) from Sardhok ...
Aug 6, 2025 · New Boselaphus (Mammalia, Bovidae) material is excavated from Pabbi Hills of Sardhok, northern Pakistan. Recently excavated bovid material ...
[25]
https://www.researchgate.net/publication/331842797_Fossils_of_Boselaphus_Bovini_Bovidae_Ruminantia_from_Sardhok_Pleistocene_of_Pakistan
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[PDF] Fossils of Boselaphus (Bovini: Bovidae - CABI Digital Library
Nov 14, 2017 · Genus Boselaphus Blainville, 1816. Type species: Boselaphus tragocamelus (Pallas 1766). Generic giagnosis: As in Pilgrim (1939). Boselaphus cf.Missing: scientific paper
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[PDF] <i>Boselaphus tragocamelus</i> (Artiodactyla: Bovidae)
Jun 6, 2008 · Abstract: Boselaphus tragocamelus (Pallas, 1766) is a bovid commonly called the nilgai or blue bull and is Asia's largest antelope.Missing: paper | Show results with:paper
[28]
Fossils of Boselaphus (Bovini: Bovidae: Ruminantia) from Sardhok ...
New Boselaphus (Mammalia, Bovidae) material is excavated from Pabbi Hills of Sardhok, northern Pakistan. Recently excavated bovid material in the Pleistocene ...
[29]
Holy yet Hindered: Habitat use patterns of a large herbivore reveal ...
The species is endemic to the Indian subcontinent, occurring naturally in India, Nepal, and Pakistan. Introduced populations are found in the United States ( ...2. Materials & Methods · 2.2. Nilgai Occurrence Data · 4.2. Nilgai Habitat Use And...
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The Reappearance of Nilgai (Boselaphus tragocamelus) in ...
Aug 2, 2024 · It is endemic to peninsular India and small parts of Pakistan and Nepal, has been extirpated from Bangladesh, and has been introduced in the ...
[31]
Habitat Suitability and Conflict Zone Mapping for the Blue Bull ...
Mar 5, 2023 · The Blue bull (Boselaphus tragocamelus), commonly known as 'Nilgai ... Population status, distribution and potential threats of the Blue Bull ...
[32]
(PDF) Boselaphus Tragocamelus (Artiodactyla: Bovidae)
Aug 9, 2025 · Boselaphus tragocamelus (Pallas, 1766) is a bovid commonly called the nilgai or blue bull and is Asia's largest antelope.
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[PDF] Social Behavior and Movement Ecology of Nilgai Antelope
Differences in behaviour of the nilgai (Boselaphus tragocamelus) during foraging in forest versus in agricultural land. Journal of Tropical. Ecology 32:469–481.Missing: speed | Show results with:speed
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[PDF] A supplementary report of the Pleistocene bovids from the Pabbi ...
The extinct ancestor of Boselaphus tragocamelus, Boselaphus namadicus, was recorded by Rütimeyer (1878) as Portax namadicus. Lydekker (1878) noted that there ...
[35]
In-Depth Look at The Rise of Nilgai Hunting in Texas
Nilgai have cemented themselves as a mainstay in the South Texas ecosystem with a growing estimated population over 50,000, which is particularly enticing when ...Missing: IUCN | Show results with:IUCN
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Nilgai Antelope In Northern Mexico As A Possible Carrier For Cattle ...
Sep 5, 2025 · However, reintroduction of native white-tailed deer (Odocoileus virginianus Zimmerman) and introduction of exotic nilgai antelope (Boselaphus ...
[37]
Understanding Exotic Species in Texas: Impact and Management
Jan 1, 1995 · Studies indicate that axis, sika, fallow, and nilgai are dominant competitors with native deer for food, and on some heavily populated ranges, ...
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Dietary Competition among Nilgai, Cattle, and Deer: What We Can ...
⦁ Nilgai compete directly with white-tailed deer. Rangeland where nilgai are present will not support as many deer as rangeland with no nilgai. ⦁ Cattle diets ...
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Antelopes go from pests to a plague in Texas - Los Angeles Times
Apr 29, 2007 · Exotic nilgai, once imported as hunting quarry and now flourishing, carry ticks that could wipe out cattle.
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Wild Cam: Fence-busting feral antelope open up Texas range
Oct 30, 2019 · While property owners can profit from allowing hunters access to the nilgai on their land, the antelopes are also a vector for disease. Nilgai ...
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Boselaphus tragocamelus (Pallas, 1766) - Plazi TreatmentBank
Nov 29, 2024 · The Nilgai is a large bovid, somewhat equine in appearance, with a deep and compressed neck, a tufted tail reachling the hocks, hindlimbs that are shorter than ...
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Discovery of large mammalian fossils at Morgaon, District Pune ...
The fossil assemblage includes nilgai (Boselaphus tragocamelus) and equine remains from Pleistocene deposits. 16 skeletal specimens were identified, mostly ...
[43]
[PDF] Food Habits of Nilgai Antelope in Texas - University of Arizona Journal
Nilgai's diet consists of 60% grasses, 25% forbs, and 15% browse, eating more browse when food is scarce. They prefer open areas with cover and water.
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Food habits of Nilgai (Boselaphus tragocamelus) in Van Vihar ...
Dec 9, 2015 · The diet of the nilgai consisted of diverse species of food plants. Nilgai in total used 123 plant species that include 37 trees, 34 herbs, 25 ...
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Understanding Human-Nilgai negative interactions in India
Jun 26, 2024 · Cereals ranked third (20%) in the most affected crop category, with wheat and maize being crop types frequently sustained nilgai raids. Oil ...
[46]
Nilgai | Habitat, Diet & Conservation - Britannica
Nilgai is the Hindustani word for “blue cow,” which describes the blue-gray of adult bulls. (Cows are orange-brown.) The nilgai's conformation, however, is more ...
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[PDF] Mineral content and wildlife use of soil licks in southwestern Nepal
Since the licks contained large amounts of sodium, and com- mon langurs were observed many times on artificial salt licks, sodium may be one of the elements ...
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THE INTENSITY OF SEXUAL SELECTION PREDICTS WEAPON ...
May 17, 2007 · This study provides evidence that both male horn length and sexual body size dimorphism are under sexual selection in the Bovidae.
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The origins of sexual dimorphism in body size in ungulates - PubMed
The hypothesis states that sexual size dimorphism evolved in a three-step process. Initially, ancestral monomorphic and monogamous ungulate species occupying ...
[50]
https://www.researchgate.net/profile/Stein_Moe/publication/238029290_Mineral_content_and_wildlife_use_of_soil_licks_in_southwestern_Nepal/links/559a58fe08ae793d13808da3/Mineral-content-and-wildlife-use-of-soil-licks-in-southwestern-Nepal.pdf
[51]
Nilgai Antelope (Boselaphus tragocamelus)
This large antelope, often called the blue buck, once occupied large regions of Pakistan, India and Nepal, but the range has recently contracted. Its first ...
[52]
Nilgai - The Farm at Walnut Creek
Nilgai have thin legs and a robust body that slopes down from the shoulder. Their long, narrow heads are topped by two small conical horns which are straight ...
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Origin, paleoecology, and paleobiogeography of early Bovini
It has long been known that Bovini originated from “Boselaphini”, a paraphyletic assemblage of early Bovinae including also the living nilgai (Boselaphus ...
[54]
Revision of the boselaphin bovid Miotragocerus monacensis ...
Jun 1, 2015 · M. monacensis can be assigned to the basal Boselaphini based on the plesiomorphic features in the dentition and characters of the postcranial ...
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[PDF] Middle Miocene bovids from Mae Moh Basin, Northern Thailand
Eotragus is the oldest bovid possessing true horn cores, and first appears in the fossil record around 18 Ma (Gentry 2000;. Solounias 2007). Some Eotragus ...
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[PDF] The fossil record and evolution of Bovidae: State of the field
This paper presents a summary of the current state of knowledge on the evolution and fossil record of Bovidae and identifies topics and problems for further ...
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Bovine satellite DNAs – a history of the evolution of complexity and ...
711b seems to have originated only after the divergence of Tragelaphini, Boselaphini and Bovini from the other tribes of the Bovinae subfamily (10–15 MY ago) ...
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DNA systematics and evolution of the artiodactyl family Bovidae.
Eotragus, which has been assigned to the tribe Boselaphini. (7). A rate of divergence for this tribe, using transversions only, was therefore calibrated ...
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(PDF) Revision of the boselaphin bovid Miotragocerus monacensis ...
Jul 18, 2015 · Revision of the boselaphin bovid Miotragocerus monacensis STROMER, 1928 (Mammalia, Bovidae) at the Middle to Late Miocene transition in Central ...
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Boselaphini | mammal tribe - Britannica
There are 143 different species and 50 genera of Bovidae, including one completely new species placed in its own genus, the saola, discovered in the 1990s in ...Missing: taxonomic | Show results with:taxonomic
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Dietary niche partitioning among fossil bovids in late Miocene C3 ...
Teeth of late Miocene bovids referred to Bovini and “Boselaphini” were subjected to enamel stable carbon and oxygen isotope analysis to test paleoecological ...
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[PDF] ruminantia) from dhok pathan formation of siwaliks
The boselaphines appear to decline in the Soan Formation. (3.4-0.6 Ma) and only one extinct species Boselaphus namadicus is recorded. ... This documents the ...
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A supplementary report of the Pleistocene bovids from the Pabbi ...
Nov 3, 2021 · The fourth upper deciduous premolar of B. namadicus is reported for the first time from the Pleistocene of the Siwalik Group. These taxa are ...
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Continuity of mammalian fauna over the last 200,000 y in the Indian ...
Boselaphus sp. +, +, —, —, +, +, +. Antilope cervicapra, +, +, —, —, +, +, —. Gazella sp. + ... The fauna of the Kurnool Caves. Palaeontologica Indica Series X.
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Observations on the Upper Siwalik Formation and Later Pleistocene ...
Siwalik history closes during the middle Pleistocene and is followed by alternating erosion and deposition of loessic beds beneath which a younger Paleolithic ...
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The taphonomic record of Upper Siwalik (Pinjor stage) landscapes ...
... fossil accumulation were large predators and streams. The fauna was dominated by herbivores (particularly bovids) >100 kg adult body size. Although buried ...
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Gujarat's Nilgai Population Leaps 117% In 10 Years
Feb 7, 2022 · The population of blue bull also known as nilgai, the largest Asian antelope, has increased in Gujarat by an alarming 117.27% in the past 10 years.<|control11|><|separator|>
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Population ecology and herd dynamics of blue bull Boselaphus ...
Jul 5, 2025 · Its population density and encounter rate ranged from 0.318 to 5.79 animals/km2 and 0.145–1.316 animals/km respectively. The herd composition ( ...Missing: diet | Show results with:diet
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[PDF] Their Population Structure, Density and Biomass in Western India
An average density of 40.52 individuals km of Nilgai ... The Chinkara occurred at only one site and its density was similar to that of Gir forest as reported by.
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[PDF] status of Tigers Co-predators & Prey in India, 2022 - NTCA
Jul 26, 2023 · India harbours approximately 75% global wild tiger population which is reflective of our conservation initiatives thoroughly supported by robust ...
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(PDF) Distribution of Blue Bull (Boselaphus tragocamelus) and its ...
Aug 1, 2025 · Open grazing, invasive species, predation by tiger and flash flood were the major threats to the blue bull as perceived by the local people.
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BEHAVIORAL DYNAMICS OF CAPTIVE BLUE BULLS (Boselaphus ...
In Pakistan, it is classified as endangered due to habitat loss, poaching, and human-wildlife conflict (IUCN, 2023). The establishment of protected areas ...
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[PDF] Distribution of Blue Bull (Boselaphus tragocamelus) and its ...
The major global threats are poaching, retaliatory killing in response to crop raiding, predation by tiger and habitat destruction (Jnawali et al., 2011), which ...
[74]
Challenges of Conserving Blue Bull (Boselaphus tragocamelus ...
... threats to nilgai including illegal hunting, poisoning, electric fences and habitat deterioration. Successful nilgai conservation will require further ...
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[PDF] Wildlife Services - usda aphis
The high reproductive rate and adaptability of feral swine has resulted in populations that have dramatically increased in size and distribution.
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Haryana's Nilgai Culling Policy Triggers Ethical and Environmental ...
Boselaphus tragocamelus. IUCN Status, Least Concern. Legal Protection, Schedule III, Wildlife Protection Act, 1972. Region Found, India, Nepal, Pakistan. State ...
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[PDF] Appendices I, II and III - CITES
May 4, 2023 · Boselaphus tragocamelus (Pakistan). Bubalus arnee {Excludes the ... Appendices. II. III. Brookesia spp. (Except the species included in Appendix I).
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Characteristics of the Nilgai - BYJU'S
In the IUCN Red List the Nilgai is categorised as 'Least Concenced In India', the Nilgai is protected under Schedule III of the Wildlife Protection Act of 1972.
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[PDF] a perceptive study on feeding pattern of nilgaii, boselaphus ...
Nilgaii are grazers, with their diet consisting of 60% grasses, 25% forbs, and 15% browse, and are considered mixed feeders.
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Nongame, Exotic, Endangered, Threatened & Protected Species
A hunting license is required for the take of nongame species. There are no closed seasons, bag limits or possession limits; and, they may be hunted at any ...
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NILGAI IN SOUTH TEXAS: NUISANCE OR ASSET?
The value of a nilgai hunt may range from $1 500 to $2 000 for a bull and $500 for a cow and the price of nilgai meat may go from $11 per lb for hamburger ...
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Ghodpara (Nilgai): Culling, Habitat, and Conservation Status - Prepp
Despite human-wildlife conflict, Nilgais hold a stable population and are legally protected under various conservation frameworks: IUCN Red List: Least Concern ...