TYPE: Commentary

Arrival, rise, fall, and again rise of the Asiatic lion Panthera leo leo in India

Hari Shanker Singh¹*

¹National Board for Wildlife, Indira Paryavaran Bhawan, Jor Bagh Road, New Delhi-110003, India

RECEIVED 03 February 2025
ACCEPTED 16 June 2025
PUBLISHED 18 June 2025

https://doi.org/10.63033/JWLS.BHUU5534

Abstract

Many animal populations have shifted their distribution and emigrated to new areas in response to climate change, and lions in India have had a similar story. This commentary examines historical records, environmental barriers, climate change in the region of Indus-Sarasvati rivers that created conditions for lions’s entry in India. Recovery of artefacts of several wild animals and near absence of lion in these ancient artworks at any site of the Indus-Sarasvati civilization revealed that lion was absent or rare beyond the east of the Indus River in India before 2000 BCE. Environment progressed from the moist conditions to dry in the Indus-Sarasvati region between 2600 and 1500 BCE, discharge of snow water from the Himalayas declined and many large rivers of Indus-Sarasvati system transformed into seasonal rivers. Also, the dense forests transformed to thorn forests and savannah like vegetation over a period after 2000 BCE. Subsequently, Asiatic lions from the ancient Persian land got suitable environment and opportunity to cross the Indus valley to the east. After a period, lion population flourished reaching its peak in India during Buddha-Mauryan-Gupta period (600 BCE – 500 AD). Hunting records revealed that the distribution and abundance of lions remained at its peak during the Mughal and early British period before end of the Little Ice Age during the mid-19ᵗʰ century. Subsequently, fall of the Asiatic lion was sharp as numbers declined below hundred. After a long critical period of ups and downs, Asiatic lion’s number and distribution range is on consistent rise since declaration of Gir Lion Sanctuary Project in the early 1970s. Since then, lion population has increased by five folds (891 individuals in 2025) along with impressive recovery of wild ungulate population, turning management approach into one of the most successful wildlife conservation stories in the world. Considering dispersion trend of the lions and their arrival to Barda forest, an identified alternative site for lion, Gujarat Government has prepared a long term comprehensive project, “The Project Lion @2047- A vision of Amrut Kal” to secure and manage the growing lion population and its potential habitats distributed in entire Saurashtra region.

Keywords: Climate change, Indus valley civilization, lion distribution, lion history, lion population.

Introduction

Before the recent genetic studies on the surviving lion populations from different regions in the world, biologists placed all African lions within a single subspecies, Panthera leo leo, and the Asian lions as the second zoo-geographic subspecies, P. leo persica (O’Brien et al., 1987; Bauer et al., 2016; Jhala et al., 2019). These two subspecies had possibly diverged around 55,000 to 200,000 years ago (O’Brien et al., 1987). However, recent studies found that the Western and Central African lions are more closely related to the Indian lions in the Gir forests than those found in South and East Africa (Bertola et al., 2022). Their work has improved the subspecies classification, categorizing the surviving lions in two subspecies: Panthera leo leo– lions of Central Africa, West Africa, and Asia (India), and Panthera leo melanochaita-Southern and Eastern African lions (De Manuel et al., 2020; Bertola et al., 2022). Recent analysis using mt-DNA from the latest and old lion samples shows that the movement of lions from north Africa into Asia started sometime around 21000 years ago and probably continued till the late Holocene, bringing fresh genetic material to the existing Asian lion populations (Barnett et al., 2014). The study found evidence of separate incursions into India from North Africa through Asia Minor. The maternal lineage of the current Asiatic lion population in Gir was found to be a part of the clade of the Northern, Western, and Central African lions (Barnett et al., 2014; Jhala et al., 2019). Research by Bertola et al. (2015), who included nuclear as well as mt-DNA markers, found that the Indian lions were a distinct genetic cluster without much admixture from African lions, plausibly evolving separately after migrating to India.

Globally many animal populations have shifted their distribution and emigrated to new areas in response to climate change, including the lions (Barnett et al., 2014). During the period of the Indus Valley civilization, climate-induced vegetation shifts in the forests of the Indus-Sarasvati landscape have been well-documented (Shaffer & Lichtenstein, 1989; Enzel et al., 1999; Staubwasser et al., 2003; Cliff, 2009; Giosan et al., 2012), but there have been no corresponding investigations of wild animal emigration/dispersal. This paper logically clarifies the issue and tries to bring new facts related to the arrival, rise, fall, and again rise of the Asiatic lion in India.

When and how the Asiatic lion crossed the western Himalayan passes and entered the Indian subcontinent has remained a matter of debate due to a lack of extensive and authentic evidence. Fossil records in Sri Lanka (Manamendra-Arachchi et al., 2005) indicate lion’s presence as early as the late Quaternary, much before the estimated arrival of both modern lions and tigers into India (Jhala et al., 2019). However, this may not be relevant in the context of the entry of the modern lion into the Indian subcontinent, unless the fossil records of Sri Lanka show links to the current lions in India.

When did the Asiatic lion enter India?

Lions perhaps first entered India from the western Himalayan passes (Singh, 2017a; Rashid & David, 1992). As the lions could not possibly enter mainland India from Asia Minor/Persia without crossing the Indus River and the Indus-Sarasvati landscape. The Indus Valley civilization, also known as the Indus-Sarasvati civilization or Harappan civilization, was a Bronze Age civilization in the north-western regions of the Indian subcontinent, lasting from 3300 BCE to 1500 BCE, with a mature phase from 2600 BCE to 1900 BCE. The seals and artefacts linked to the Indus Valley civilization often depict common animals like tiger, elephant, rhino, bull, antelope, crocodile, etc., but lion is conspicuously absent (Divyabha­nusinh, 2008; Dutt et al., 2018). Except at sites of Mehrgarh civilization in the west of the Indus River, lion art was not found on seals, pottery, and terracotta at any of the hundreds of Indus Valley civilization’s sites such as Mohenjo-daro, Dholavira, Harappa, Lothal, and Rakhigarhi (Singh, 2017a). A rare figurine of a two-headed lion-like creature was recovered from an Indus Valley site, but it was likely an imported artefact (Divyabhanusinh, 2008; Singh, 2017a).

Surprisingly, lion painting or art on terracotta vessels were recovered at Mehrgarh (Baluchistan) in the west of the river Indus, one of the most important Neolithic (7000 BCE to 1700 BCE) sites in archaeology (Figure 1). The presence of lion in artefacts at the Mehrgarh in the west of the Indus River indicates a close association of the lion with people of western Indus Valley as well as people of the ancient Persian lands. Lions have had a close relationship with humans, as it was reflected in human art and culture wherever both shared the same landscapes. What was the reason for the presence of such artefacts at Mehrgarh and their absence at any other site east of the Indus River? Perhaps, dense moist forest and the mighty Indus River worked as a barrier against the movement of the lions to the east of the Indus River before the progress of aridity in the region. Therefore, the lion was plausibly not present in the east of the Indus River during the mature phase of the Indus Valley civilization.

To understand the period of lions crossing the Indus-Sarasvati region to its east, it is necessary to discuss climate, forests, and conditions of the rivers and environment. The study by Dutt et al. (2018) indicates an interval of warm and wet climatic conditions before and during mature phase of the Indus valley

sites in archaeology

Figure 1. Artefacts depicting a lion-like figure recovered from Mehrgarh, a western Indus Valley civilization site. Source: Singh (2017a) and Google images

civilization with a trend towards deteriorating climate, leading to a protracted period of cold and arid phase from 2350 to 1450 BCE in the Indus-Sarasvati region. During this protracted dry spell, the Indian summer monsoon weakened depleting the water resources in the region that likely had triggered the gradual desertion of the Harappan cities. Study suggests a more than 900 years long event of dry spell in the North-West Himalayas beginning at about 2350 BCE, which decreased the precipitation and snow melting in the Northwest Himalayas and subsequently the discharge in the Indus River system (Dutt et al., 2018). A shift in temperatures and weather patterns over the Indus valley at the beginning of 2500 BCE caused summer monsoon rains to gradually dry up, making agriculture difficult or even impossible close to Harappan cities (Giosan et al., 2012).

Shaffer & Lichtenstein (1989) envisaged a wet climate during early Harappan times, placing the mature phase in an already marked trend of aridity. The planktonic oxygen isotope ratios of the Indus delta were examined by Staubwasser et al. (2003). According to their research, the climate changed over the past 6,000 years, with the most notable shift occurring during the last phase of the fully developed Indus Valley civilization with a reduction in water flow in the Indus River. They observed that the 2200 BCE event aligns with the drying rivers and sudden decline of urban Harappan civilization in the Indus Valley (Enzel et al., 1999; Staubwasser et al., 2003; Cliff, 2009). Gupta et al. (2006) collected studies on the monsoon and other climate factors from a variety of sources, including their own, and came to the conclusion that the arid phase in the Indian subcontinent started during the mature phase of the Indus Valley civilization, coinciding with a stepwise weakening of the south-west monsoon. The arid phase might have intensified during 2000 – 1500 BCE, as has been in the Himalayas, western peninsula, and north-western India. Danino (2016) also wrote that the early mature Indus Valley civilization was the time when, in the east of the Indus system, the mighty Sarasvati dwindled to a minor seasonal river. Evidence in these scientific publications suggest that the savannah habitats existing today in western India are young.

Based on abovementioned studies, it appears to be well recognized now that climatic and environmental disruptions were a significant factor in the decline of moistness of environment, transformation of many mighty rivers into seasonal rivers, gradual change of high forests into thorn forests and savannah like vegetation, leading to final break-up of the Indus civilization. In other words, early phase or at the beginning of the mature phase of the Indus Valley civilization, the environment of the Indus-Sarasvati region was wet with dense, moist to semi-moist tropical forest, and large rivers acting as a barrier against the movement of the lion to the east of the Indus River. When many rivers of the Indus-Sarasvati system changed seasonal, discharge of snow water from the Himalayas declined in the rivers and dense forests transformed to thorn forests or savannah like vegetation over a period, lions from the Persian region in the west of the Indus River got suitable environment and opportunity at some time, most likely after 2200 BCE, to cross the Indus valley to its eastern lands in India.

There could be two reasons for the presence of lion artifacts at the site of the Mehrgarh. First, it was an extension of the Persian civilization where the lion was the dominant big cat. Second, perhaps lions crossed the Bolan pass (near Quetta) and lived in the region west of the Indus River, indicating continuity of lion distribution from the ancient Persian land to Mehrgarh through the Bolan Pass. However, the dense, moist forest in the lower Indus Valley and a bigger Indus River could have acted as an environmental barrier for the lion.

M. A. Rashid and Reuben David, the two known lion experts from India, write that the lion migrated to India through the north-western passes much before 6000 BCE (Rashid & David, 1992), but no authentic evidence was mentioned to substantiate it. The Rigveda, one of the oldest texts in the world (approximately 1700–1100 BCE), places great importance on lions, bulls/cows, and horses. The early part of the Rigveda does not mention the tiger, but the lion is mentioned, perhaps because the bulk of this text originated either in the north-western Himalayas or northern Persia, which had lions. Subsequently, the lion in culture was carried on by the people in the Rigvedic land to the Indian subcontinent.

A fresh controversy emerged after the publication of a book, “Exotic Aliens: The Lion & the Cheetah in India” (Thapar et al., 2013). The statement in the book about the Asiatic lion being introduced in India by humans was contested by wildlife experts like Divyabhanusinh and Ranjitsinh (Jhala et al., 2019). Such views of anthropogenic introduction of the Asiatic lion, opined by some naturalists and historians, have been negated by genetic studies. O’Brien (2013) writes that no African genetic lineage was ever discovered among the wild Gir lions sampled. A comprehensive analysis of molecular phylogeny indicated a clear genetic distance between the present Gir and African lion populations, indicating separate populations for a long period and no evidence of gene flow between African and Asiatic lions after the Holocene (Barnett et al., 2014; De Manuel et al., 2020). This curtails the scope for continuing discussion on the points raised by Thapar et al. (2013).

Rise of lions – Abundance and historical range in India

The lion came to India at a time when tigers and leopards had already settled in the subcontinent (Rashid & David, 1992; Bernett et al., 2014). Perhaps the lion was present in the areas inhabited by the new human migrants (Aryans) and on the route through which they migrated to India. The close proximity of lion may explain its prominence in their art and literature and the absence of tiger. It is also likely that the lions followed their prey, the livestock, and fresh genetic pool of lions entered in mainland India following human migrants and their cattle and thus began to colonize the northern and western parts of the country. Within a millennium, they expanded their habitat up to the rivers Narmada and Ganga. It may be inferred that the lion’s history in the Indian subcontinent may not be more than 4,000 years old. Kailash Sankhala, in his book “Tiger! The Story of the Indian Tiger”, says that the tiger seems to have lost its supremacy in India for some time after about 1500 BCE (Sankhala, 1978). The lion was frequently mentioned in religious and cultural works, including the Rigveda, Buddhist Jatakas stories, Panchatantra, and Sanskrit literature in general. Lions guarded the gates of the majority of temples of the ancient and early medieval periods. In the late medieval period, the lion dominated human culture in north-west India. Following Independence, the lion capital of the Ashoka pillar (300 BCE) was chosen as India’s emblem and, subsequently, its national animal. The lion, which had dominated India for over 3,000 years, was replaced by the tiger as the country’s national animal only in 1972–1973.

The Rigveda has more than fifteen references to the lion (Simha). Aryan devatasRudra and Agni- are compared to a lion in the Vedas. In Dev-Asur yug, Narsimha, a Lord Vishnu incarnation or avatar who is half man and half lion and killed Hiranyakasyipu, a great Asur king, thereby restoring Dharma. In the early Rigvedic period, before the epic age (earlier than 600 BCE), Bharat, son of the great king Dushyant and Shakuntala, played fearlessly with lion cubs in a sage’s ashram, a testament to his bravery. In the war of the ten kings, Indra, king of the Devas, provided aid to Rigvedic king Sudas of the Bharat clan against the vast host of enemies. In this war, the defeat of ten kings is compared to the defeat of a lioness by a ram. There is an interesting story in both the Ramayana and Raghuvamsa of a lion attempting to kill the cow, Nandini, who was saved by king Dilip, an ancestor of Lord Rama. These mythological stories indicate the presence of lions in the lands of their kingdoms.

It is likely that the period from the entry of the lion in India to 600 BCE saw an increase in the lion’s distribution range in the north-western and north-eastern India up to Bihar – Bengal and up to the Narmada River in the south. The Harappan civilization used the bull, the elephant, the rhino, and the tiger as cultural symbols. In Asia Minor, Europe, and Egypt, every god, goddess, and king was ‘lionized’. But in India, before Mahavir (540 BCE to 468 BCE), none of the twenty-three Tirthankars of Jainism had the lion as their symbol. Every one of them selected plants and animals, including snakes, as their symbol. King of Ikshwaku Vansh of Simhpur (now Sarnath, near Varanasi) was the father of the 11th Tirthankar, Lord Shreyansnath. The name of the kingdom, Simhpur, suggests that the lion perhaps existed when Lord Shreyansnath was born in the city (Singh, 2017a).

Queen Trishala, mother of Vardhman, who was later named as Lord Mahavir, dreamt fourteen beautiful and auspicious events after conception, including a magnificent lion, at midnight. The dream was interpreted that her son would be as powerful as a lion. He would be fearless, mighty, and capable of ruling the entire world. The lion became a symbol of Tirthankar Mahavir, but this royal animal could not become a symbol of other Tirthankars before him.

Siddhartha, who was later called Gautam Buddha after achieving enlightenment around 524 BCE, was born around 560 BCE to Sakya chieftain. He was also known as Sakyasimha, the lion of the Sakya tribe. His first sermon at Sarnath is known as Simhanad, the roar of a lion, since his voice was as loud and effective as the roar of a lion. While other gods and kings in ancient India were symbolized with animals like elephant and bull, the lion was chosen as a symbol for Gautam Buddha, Mahavir Jain, and the Mauryans. The domination of the lion in culture during the time of the Mahavir Jain, Buddha, and the Mauryas around the sixth century BCE indicates that the distribution and population of the lion reached a peak during those periods. By the time Jainism and Buddhism grew in significance, lions had a well-established distribution range in India.

During the time of Buddha, the Asiatic lion roamed from Sindh in the west to Bengal in the east. The Himalayan foot-hills and the Ganga plains formed their northern and eastern limit, while the Narmada seemed to be its southern boundary, although unconfirmed report of the lion south of Narmada River have also emerged (Rashid & David, 1992; Singh, 2017a). A question might be raised as why the lions did not reach the eastern plains of the Ganga River. There could be two reasons: first, the lions could not get enough time to cross the mighty Ganga in the plains, and the second reason could be that the dense, moist forests and Ganga river acted as an environmental barrier. Li­ons were abundant in the Gupta period (300 AD to 600 AD), as kings of the period – Chandragupta II and Kumargupta hunted lions and minted lion coins. There are plenty of records of lion hunts during the Mughal period and early British period (mid-15ᵗʰ century to 19ᵗʰ century AD). Thus, the Asiatic lion flourished from the first millennium BCE to the mid-19ᵗʰ century AD in the north-west of India.

Fall after the Little Ice Age – A critical period a hundred years ago

The Asiatic lions are now restricted to a single population in and around the Gir forests in Gujarat State of India. Due to un-restricted hunting and habitat loss, lions were extirpated from Asia Minor and a major part of the habitats in India by the end of the 19ᵗʰ century (Figure 2; Singh, 2017a).

Lion distribution in India and hunting records

Figure 2. Lion distribution in India and hunting records by the mid-19ᵗʰ century. Sources: Joslin (1973), Rashid & David (1992), Divyabhanusinh (2008)

Except for lion hunting records of the Mughal and British period, the history of lions in India before the 19ᵗʰ century, especially beyond the Gir forest, is poorly documented. Kazmi (2021) has described chronological records of lion hunts in Hurrianah (now Haryana) and the region around Delhi. His research established the existence of a good number of lions, and their large-scale hunting, which involved wiping out entire prides, including the killing or capturing of lion cubs in Haryana, leading to their extinction in a short period. He listed 26 references dealing with lions in the Haryana landscape between the years 1809 to 1823. The records also provide very interesting information on the ecology of lions in Haryana. These references mention sightings of a total of 129 to 141 lions, out of which at least 109 were conclusively killed by the hunters. Of them, as many as 80 lions were killed in a mere five-year period (1810–1815). The British record of June 18, 1811, mentioned that the lions were very plentiful in the area around Hansi, Haryana, where tigers and leopards also occurred. Kazmi (2021) also mentioned the travel diaries of Maria Nugent, the wife of Sir George Nugent, the Commander-in-Chief of India (1811–1813), who kept record of lion hunts. William Fraser, who was part of Lady Nugent’s camp, hunted plenty of lions in Haryana, mostly on foot or on horseback. Divyabhanusinh (2008) also mentions that the British encountered plenty of lions in the region around Delhi. Naturalists Richard Lydekker and J. G. Dollman claimed that Colonel George Acland Smith hunted 300 Indian lions during his stay in India (out of which about 50 lions were killed in the Delhi district) in the years leading up to and just after 1857, although this statement was contested by some naturalists (Thapar et al., 2013; Kazmi, 2021) . Available records indicate that population and distribution range declined drastically after the First War of Independence (1857) in India, which was also the ending phase of the Little Ice Age (1300 AD to 1850 AD).

During the Little Ice Age, from the early 14ᵗʰ century through the mid-19ᵗʰ century, when mountain glaciers expanded, the climate was cooler than the present days. Subsequently, habitat loss and rampant hunting, combined with the impact of a hot environment, perhaps caused a drastic decline in the distribution range and population of lions.

Available records suggest that lions occurred in almost all parts of Saurashtra during the Mughal period, and lion hunts were recorded in all districts of the region (Gee, 1964; Rashid & David, 1992; Singh, 2017a). By the 1880s, lions were restricted in and around the Barda and Alech hills, Mitiyala, Girnar, and Gir forests in the Saurashtra peninsula of Gujarat (Jhala et al., 2019). Subsequently, lions were extirpated from Barda and Alech hills and were restricted to Gir forest and adjoining areas. Since 1880, there has been an increased concern about the falling numbers of the Asiatic lion in the Gir forest. The last three decades of the 19ᵗʰ century and the first two decades of the 20ᵗʰ century were the worst for the Indian lion as its population was estimated below a hundred individuals, with few naturalists mentioning a population below 50 individuals (Gee 1964; Rashid & David, 1992). Although most of these estimates were opinions rather than a result of systematic surveys. At the beginning of the 20ᵗʰ century, Lord Curzon declined a lion hunt in the Gir forests in Junagadh because the Asiatic lion was on the verge of extinction and instead asked the Nawab of Junagadh to protect the animal. According to Fenton (1909), a British field officer, five or six lions were shot annually within Gir and about eight in the outlying areas, despite protection. The Chief Forest Officer of Junagadh mentions in 1913 that a good number of lions were hunted annually in Mitiyala, a block of forest governed by the princely Estate of Bhavnagar (Anon., 1975; Singh, 2017b).

The Statistical Account of Nawab (ex-ruler) of Junagadh in 1884, through a statement of a Britisher, Col. J. W. Watson, mentioned that the Gir lion population was about a dozen (Divyabhanusinh, 2008; Singh, 2017a). After a tour of the Gir forest in 1913, Wallinger, the Chief Forest Officer, raised alarm about a low number of lions and reported that there were no more than 20 lions in the Gir forest of Junagadh State (Anon., 1975; Singh, 2017a). However, during the same period, Major H. G. Carnagie, in 1905, mentioned about 80 surviving lions (at least 70 lions), and J. R. Ratanagar and Sir P. R. Gadell stated about 50-100 free-ranging lions in the Gir forests in 1920 (Anon., 1975). The scientific discourses later picked the lower number (a dozen to 20 individuals) as a reference of lion population at the beginning of the 20ᵗʰ century for unknown reasons, ignoring observations of other naturalists. At that time, lions occurred in Rajkot and Bhavnagar states beyond the boundaries of the Gir forests governed by the Nawab of Junagadh. During the first census of the Asiatic lion in 1936 in and around the Gir forests, 287 individual lions were counted (Anon., 1975; Singh, 2017b). If the lion number was below two dozens in 1913, how could it reach 287 individuals at the time of the first lion census in 1936? It is evident that the free-ranging lion population most probably never dropped below fifty during the entire history of the Indian lion. Ignoring several reports and logical findings, and sticking to two individual statements, numerous scientific papers and books have been published discussing and ascertaining an unrealistic population rebound. These scientific publications have created an improbable narrative that the present population of the Gir lion is built up from a dozen lions that survived in the Gir forests. These narratives need recalibration to correct the distorted recent history, and future discourses must be more careful.

Accurate information about the distribution range and abundance of lions in India before the first lion census in 1936 in the Gir forest is not well understood. However, it is evident that the distribution range and population of lions reached their lowest level at the end of the 19ᵗʰ century and the beginning of the 20ᵗʰ century. At present, the Gir forest is synonymous with the Asiatic lion.

Rising again – Recovery from the verge of extinction

Along with Asiatic lions, other carnivores such as the Indian leopard (Panthera pardus fusca), striped hyena (Hyaena hyaena), Indian golden jackal (Canis aureus), jungle cat (Felis chaus), Indian fox (Vulpes benghalensis), honey badger (Mellivora capensis), and rusty-spotted cat (Prionailurus rubiginosus) occur in the Gir landscape. Nine wild prey species, consisting of seven wild ungulates (spotted deer Axis axis, sambar Rusa unicolor, blue bull Boselaphus tragocamelus, four-horned antelope Tetracerus quardricornis, blackbuck Antelope cervicapra, Indian gazelle Gazella benneti, and wild pig Sus scrofa), one primate (hanuman langur Semnopithecus entellus), and one large bird (Indian peafowl Pavo cristatus) are the main wild food for major carnivores in the Gir forest (Joslin, 1973; Anon., 1975). Lions are primarily dependent on wild ungulates like spotted deer, blue bull, sambhar and wild pigs, and domestic animals like cattle, buffalo, whereas leopard’s dependency is high on spotted deer, sambar, wild boar, Hanuman langur, peacock, Indian hare and medium to small domestic animals (Anon., 1975; Singh, 2017b). Beyond Gir boundaries, lions’ dependency for food is high on cattle, buffalo, blue bull, wild pig, and carcasses of domestic animals (Jhala et al., 2019).

The Gir Lion Sanctuary was created in 1965, and the Gir Lion Sanctuary Project was launched in 1972. Before the implementation of the Gir Lion Sanctuary Project, the prey population was very low, and lions’ dependency was mainly on domestic livestock-cattle and buffalo (Joslin, 1973; Anon., 1975). Implementation of the project was one of the most successful conservation stories, as the lion population consistently increased by five folds (Figure 3) and seven species of wild ungulates also increased by many folds (5,600 in 1973 to 91,300 in 2019) during the last five decades (Joslin, 1973; Anon., 1975; Singh, 2017b; Jhala et al., 2019; Ram et al., 2023a).

At present, lion is found in seven districts– major parts of Junagadh, Gir-Somanath, Amreli, and Bhavnagar, some parts of Rajkot, Porbandar, Devbhumi, Dwarka, and occasional visits in Surendranagar district. In the winter of 2023, one lion and two lionesses arrived in the Porbandar district and settled in the Barda forest, a site identified in the 1979 as an alternative habitat for lions. Subsequently, they were joined by three more lionesses. Breeding of these lions was documented during the last two and a half years. At present, there are 17 individual free-ranging lions (one male lion, 5 lionesses, 2 sub-adults over one year, and 9 cubs) in the Barda forest. Thus, the restoration and development of an alternate lion population away from the Gir landscape is currently in progress at Barda forest.

In 2015, lion counting was done in about 22,000 km² , which was further increased to about 30,000 sq. km in 2020, and 35,000 km² in 2025 due to continued dispersion of the lion (Gujarat

Growth of lion population

Figure 3. Growth of lion population in the Gir landscape over the last century.

Forest Department, 2025). The population of a minimum of 891 lions at 358 locations in 2025 was female-biased (330 females, 196 males, 140 sub-adults, and 225 cubs). The network of Gir Protected Areas (Gir National Park, Gir Wildlife Sanctuary and Paniya Wildlife Sanctuary), held the largest number of lions (n= 394) as source/core population, followed by seven satellite populations and one alternate population: (i) Savarkundla-Liliya and adjoining areas (n= 125), (ii) the south-eastern coast in Amreli district (n= 94), (iii) Bhavnagar mainland-Hippavadli Zone (n= 103), (iv) Girnar Wildlife Sanctuary and adjoining area (n= 54), (v) the south-western coast in Gir–Somnath district (n= 25), (vi) the coast of Bhavnagar district (n= 15), (vii) Mitiyala Wildlife Sanctuary and adjoining area (n= 32), (viii) Barda Wildlife Sanctuary – an alternative site (n= 17), and corridors and other areas (n= 32) (Figure 4; Gujarat Forest Department, 2025). As per the lion census in May 2025, more lions are outside the Gir forest boundaries (55.8%) than the lions within it (44.2%; Gujarat Forest Department, 2025). Naturalists working in and around the Gir forests have a strong opinion that the lion population is growing consistently at the same or a higher rate. If the compound annual growth (5.74 %) of the last five years persists in the lion conservation landscape, the population may continue to disperse in the entire Saurashtra region covering 11 districts. Increasing population and detection of death cases during the last ten years also reveal a high population of lions (Ram et al., 2023b). The Forest Department reported in the Gujarat Legislative Assembly that 313 lions died in two years – 2019 and 2020 (154 deaths took place in 2019 and 159 in 2020, comprising 90 lionesses, 71 male lions and 152 cubs) and average annual deaths of lions during the last ten years, from 2015 to 2024, was 122 individuals (pers. comm. Gujarat Forest Department).

The lion count revealed that the lion number within the Gir Protected Area has marginally increased, but the rate of increase was consistently high beyond the boundaries of the Gir forests in the satellite areas. The lion distribution range has expanded from about 10,000 sq. km. in 1995 to about 35,000 sq. km. in 2025 (Singh, 2017b; Ram et al., 2023b; Gujarat Forest Department, 2025). Also, lion densities in Girnar, Savarkundla-Liliya, and the coastal zone of Amreli are higher than the densities in the best lion habitat within the Gir Protected Area. The recovery of the big cat in the lion conservation landscape in the Saurashtra region is a success story, accrediting the efforts of the local community and the Gujarat Government. This story of lions and the people living in proximity in the Saurashtra region of Gujarat provides the world with a plausible model of carnivore co-existence (Singh, 2017b; Jhala et al., 2019).

Emerging future scenario

Establishing a second population of free-ranging lions without dependency on the Gir forest has been the most important conservation priority. Kuno National Park in Madhya Pradesh was identified and developed as an alternative site for the Asiatic lion, but lions could not be translocated till date due to socio-political reasons, despite the direction of the Supreme Court of India. Barda forest in Porbandar and Jamnagar districts was also identified as an alternative habitat within Gujarat by the State Government in 1979, but the lion populations could not be restored there as well. Migration of a lion and five lionesses in the Barda forests (quite distant from the source population in the Gir forest) and the successful birth of litters of all five lionesses in recent years has moved the progress in the right direction. The State Forest Department has initiated several activities to restock Barda and the surrounding area with lions and wild ungulates. Recent development indicates that, in a few years, Barda forest is expected to be fully rehabilitated by the lions, building an alternative site.

358 locations across Gujarat(Lion)

Figure 4. During the latest Asiatic lion census on 12-13 May 2025, a total of 891 lions were recorded at 358 locations across Gujarat,India. Source: Gujarat Forest Department 2025

When the population reached a saturation level in and around the Gir forests in the 1990s, the lions gradually dispersed to new areas where they had occurred a century ago. The distribution range of the Asiatic lion was doubled in the four districts (Jhala et al., 2019) in three decades, which further expanded to seven districts in 2025 (Gujarat Forest Department, 2025). Conservation measures beyond Protected Areas were extended to the entire lion distribution range. The Gujarat Forest Department reports that the free-ranging population of Asiatic lions is widespread in the multi-use landscape of the Saurashtra region. In several villages, forest patches, wastelands, and community lands are available for resting and sheltering, although not all of them may be suitable sites for lion breeding. Considering the dispersion trend of the lions, the Gujarat Forest Department has prepared a long -term comprehensive project, “The Project Lion @2047- A vision of Amrut Kal”, to secure and manage the growing lion population and its habitats (Gujarat Forest Department, 2023a). This project, covering Wildlife Sanctuaries and National Parks, Reserved Forests, Protected forests, Un-classified forests, Reserve Vidis (Reserved Grasslands), and Non-Reserve Vidis in 11 districts of Saurashtra in Gujarat State, has been approved by the National Board for Wild Life in its 7th meeting at Gir-Sasan held on 3rd March 2025.

With the increasing lion population, the likelihood of negative human-lion interaction is also increasing. However, a high density of the Indian leopard in the lion distribution range is also a big cause of man-wildlife conflict (Vasavada et al., 2020). In 2023, a total of 995 leopards were counted in the lion conservation landscape in the four districts, viz., Junagadh, Amreli, Gir Sasan-Somnath, and Bhavnagar districts (Gujarat Forest Department, 2023b). Thus, over 1,880 big cats with high density occur in the Lion Conservation Landscape. Wild prey biomass within the Gir forests is reasonably high. Due to a high concentration of big cats, human-big cat conflicts are bound to occur. As per the data provided by the Gujarat Forest Department, average annual human deaths due to lion attacks during the last ten years (2015-24) were 3.5 per year, with average annual attacks of 18.9 per year. In comparison, average human deaths due to leopard attacks were 10.5 per year, with average annual attacks of 62 per year during the same period. During the last five years (2020-24), compensation was paid for 28,798 livestock kills (average annual kills- 5760 livestock per year) by lions and leopards, and over 80 % of those were by the lions. There were rare intentional attacks by lions on human beings, as most of the attacks were due to human errors or mischiefs. Payment of adequate compensation plays a crucial role in mitigating conflicts, but leopard attack on human beings are a major cause of human-wildlife conflicts. As lions kill blule bulls and wild boars and protect crops from their raids, people honor this majestic cat and accept its presence in the villages.

Beyond Gir Protected Areas, the population of blue bull and wild pig is good, but the other five ungulates are also present with low density. As per wildlife counting in 2023 by the Gujarat Forest Department, there were 155,360 wild ungulates outside the Gir Protected Area in 11 districts of Saurashtra, but the population of unguarded stray/feral cattle was much more than the wild ungulates. The dependency of the lions for food beyond the Gir forest boundaries is mostly on blue bull, wild boar, stray and domestic cattle, buffalo, and carcasses of the domestic animals (Jhala et al., 2019; Vasavada et al., 2020). The prevailing situation in this human-dominated multi-use landscape beyond the boundaries of Gir Protected Area indicates the scope of successful lion dispersion in other districts of the Saurashtra region in the future.

Conclusion

Recovery of artefacts of several wild animals and absence of any such craft of lion at sites of the Indus-Sarasvati civilization revealed that the lion was absent or rare in India beyond the east of the Indus River during the mature age of the civilization. Subsequently, when many rivers of Indus-Sarasvati system turned seasonal, discharge of snow water from the Himalayas declined and dense forests transformed to thorn forests or savannah like vegetation over a period, lions from the ancient Persian lands in the west of the Indus River got suitable environment and opportunity at some times after 2200 BCE to cross the Indus valley to its eastern lands and entered the Indian subcontinent, bringing major influx of lion population in India. After entering the Indian subcontinent from the Persian region, the lions population reached its peak during the Buddha period (6th century BCE). Lion distribution and abundance remained at the peak during the Mauryan and Gupta period (320 BCE to 550 AD). Hunting records reveal that the lion’s distribution range was in the entire North-West India, south of the Ganga River and North of the Narmada River. The records also reveal that, during the first half of the 19th century AD, lions were abundant in Haryana, the region around Delhi, Rajasthan, and a part of Central India and Gujarat.

The period of the Little-Ice Age was over by 1850. Climate change, loss of habitats, and rampant hunting decimated lions from their entire distribution range. By 1880, lions were confined to the Gir-Girnar-Mitiyala landscape with a low number. As per the observation of naturalists and hunting records, the Asiatic lion population was at its lowest range during the period from 1880 to 1920, but several other observations and records, including the Lion census in 1936 (287 individual lions), indicate that the population of lions never dropped below 50 individuals. Subsequent scientific publications have followed a misleading notion that the present population of the Gir lion was built up from a dozen lions that survived in the Gir forests. Prevailing dispersion trend and future vision and plan of the Gujarat State in India for lion conservation may lead to consistent recovery of lion population and expansion of distribution range in the entire Saurashtra region beyond the present Gir Lion Conservation Landscape.

Acknowledgement

I thank Chief Wildlife Warden of Gujarat State and other forest officials for inviting me as observer for sixteenth lion census and providing me necessary data and information.

CONFLICT OF INTEREST
The author declares no conflict of interest.

DATA AVAILABILITY
No additional data was used in this research.

Edited By
Vishnupriya Kolipakam
Wildlife Institute of India, Dehradun, India.

*CORRESPONDENCE
Hari Shanker Singh
hss.ifs@gmail.com

CITATION
Singh, H. S. (2025). Arrival, rise, fall, and again rise of the Asiatic lion Panthera leo leo in India. Journal of Wildlife Science, 2(2), 37-45. https://doi.org/10.63033/JWLS.BHUU5534

COPYRIGHT
© 2025 Singh. This is an open-access article, immediately and freely available to read, download, and share. The information contained in this article is distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), allowing for unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited in accordance with accepted academic practice. Copyright is retained by the author(s).

PUBLISHED BY
Wildlife Institute of India, Dehradun, 248 001 INDIA

PUBLISHER'S NOTE
The Publisher, Journal of Wildlife Science or Editors cannot be held responsible for any errors or consequences arising from the use of the information contained in this article. All claims expressed in this article are solely those of the author(s) and do not necessarily represent those of their affiliated organisations or those of the publisher, the editors and the reviewers. Any product that may be evaluated or used in this article or claim made by its manufacturer is not guaranteed or endorsed by the publisher.

Anon. (1975). The Gir Lion Sanctuary Project. Government of Gu­jarat. p.131.

Barnett, R., Yamaguchi, N., Shapiro, B., Ho, S. Y. W., Barnes, I., Sa­bin, R., Werdelin, L., Cuisin, J., Larson, G. (2014). Revealing the maternal demographic history of Panthera leo using ancient DNA and a spatially explicit genealogical analysis. BMC Ecology and Evolution, 14, 70. https://doi.org/10.1186/1471-2148-14-70

Bauer, H., Packer, C., Funston, P. F., Henschel, P. & Nowell, K. (2016). Panthera leo. The IUCN Red List of Threatened Species. http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T15951A107265605.en

Bertola, L. D., Vermaat, M., Lesilau, F., Chege, M., Tumenta, P. N., Sogbohossou, E. A., Schaap, O. D., Bauer, H., Patterson, B. D. et al. (2022). Whole genome sequencing and the application of a SNP panel reveal primary evolutionary lineages and genomic variation in the lion (Panthera leo). BMC Genomics, 23(1), 321. https://doi.org/10.1186/s12864-022-08510-y

Bertola, L. D., Tensen, L., van Hooft, P., White, P. A., Driscoll, C. A., Henschel, P., Caragiulo, A., Freedman, I. D., Sogbohossou, E. A., et al. (2015). Autosomal and mtDNA Markers Affirm the Dis­tinctiveness of Lions in West and Central Africa. PLoS ONE,11(3), e0149059. https://doi.org/10.1371/journal.pone.0137975

Clift, P. (2009). Harappan Collapse. Geoscientist, 19(9), 18–22.

Danino, M. (2016). Environmental factors in the decline of the Indus – Saraswati Civilisation. In: The Environment and Indi­an History. The C. P. Ramaswamy Aiyar Foundation, Chennai. pp.132-48.

De Manuel, M., Barnett, R., Sandoval-Velasco, M., Yamaguchi, N., Vieira, G. F., Mendoza, Z. M. L., Liu, S., Martin, M. D., Sinding, S. H. M. et al. (2020). The evolutionary history of extinct and living lions. Proceedings of the National Academy of Sciences, 117(20), 10927–10934. https://doi.org/10.1073/pnas.1919423117

Divyabhanusinh (2008). The lion of India. Black kite. Pauls Press, New Delhi.

Dutt, S., Gupta, A. K., Wünnemann, B., Yan, D. (2018). A long arid interlude in the Indian summer monsoon during ∼4,350 to 3,450 cal. yr BP contemporaneous to displacement of the Indus valley civilization. Quaternary International, 482, 83-92. https://doi.org/10.1016/j.quaint.2018.04.005

Enzel, Y., Ely. L. L., Mishra, S., Ramesh, R., Amit, R., Lazar, B., Rajag­uru, S. N., Baker, V. R. & Sandler, A. (1999). High-Resolution Holo­cene Environmental Changes in the Thar Desert, Northwestern India. Science, 284(5411), 125–28. https://doi.org/10.1126/science.284.5411.125

Fenton, L. L. (1909). The Kathiawar Lion. Journal of the Bombay Natural History Society, 19, 4-15.

Giosan, L., Clift, P. D., Macklin, M. G., Fuller, D. Q., Constantines­cu, S., Durcan, J. A., Stevens, T., Duller, G. A. T., Tabrez, A. R. et al. (2012). Fluvial Landscapes of Harappan civilization. Proceedings of the National Academy of Sciences, 109(26), E1688– E1694. https://doi.org/10.1073/pnas.1112743109

Gupta, A. K., Anderson, D. M., Pandey, D. N. & Singhvi, A. K. (2006). Adaptation and human migration, and evidence of agriculture coincident with changes in the Indian summer monsoon during the Holocene. Current Science, 90(8), 1082–1090. https://www.jstor.org/stable/24089270

Gee, E. P. (1964). The Wildlife of India. Collins, James Place Lon­don. p.192.

Gujarat Forest Department (2023a). The Project Lion @2047-A vision of Amrut Kal. Gujarat Forest Department, Gandhinagar, Gujarat.

Gujarat Forest Department (2023b). Leopard counting report. Gujarat Forest Department, Gandhinagar, Gujarat.

Gujarat Forest Department (2025). Report on 16th Lion Popula­tion Estimate. Gujarat Forest Department, Gandhinagar, Gujarat. p.36

Jhala, Y. V, Banerjee, K., Chakrabarti, S., Basu, P., Singh, K., Dave, C. & Gogoi, K. (2019). Asiatic Lion: Ecology, Economics, and Pol­itics of Conservation. Frontiers in Ecology and Evolution, 7, 312. https://doi.org/10.3389/fevo.2019.00312

Joslin, P. (1973). The Asiatic lion: a study of ecology and behavior. Ph.D. thesis. Department of Forestry and Natural Resources, Uni­versity of Edinburgh, UK, p.249.

Kazmi, R. (2021). The Last Hurrah of The Hurrianahâ Lion Pan­thera Leo Persica. Part I: A Chronological Record of Hitherto Un­known References to Lion Hunting in Present-Day Haryana and Related Reports. Journal of the Bombay Natural History Society, 118. https://doi.org/10.17087/jbnhs/2021/v118/156545

Manamendra-Arachchi, K., Pethiyagoda, R., Dissanayake, R., & Meegaskumbura, M. (2005). A second extinct big cat from the late quaternary of Sri Lanka. The Raffles Bulletin of Zoology, 12, 423–434.

O'Brien, S. J., Joslin, P., Smith III, G. L., Wolfe, R., Schaffer, N., Heath, E., Ott‐Joslin, J., Rawal, P. P., Bhattacharjee, K. K. & Martenson, J. S. (1987). Evidence for African origins of founders of the Asiatic lion species survival plan. Zoo Biology, 6(2), 99–116. https://doi.org/10.1002/zoo.1430060202

Ram, M., Sahu, A, Srivastava, N, Chaudhary, R, Jhala, L, Zala, Y. (2023a). The semi-arid ecosystem of Asiatic Lion Landscape in Saurashtra, Gujarat: Population density, biomass and conser­vation of nine wild prey species. PLoS ONE, 18(9), e0292048. https://doi.org/10.1371/journal.pone.0292048

Ram, M., Vasavada, D., Tikadar, S., Jhala, L. & Zala, Y. (2023b). Population status and distribution of endangered Asiatic lions in Gujarat, India. European Journal of Wildlife Research. 69(87). https://doi.org/10.1007/s10344-023-01720-z

Rashid M. A. & David, R. (1992). The Asiatic Lion. Department of Environment, Government of India.

Sankhala, K. (1978). Tiger! the story of the Indian tiger. Collins, London. p. 220

Singh, H. S. (2017a). The Asiatic lion-the Pride of Gujarat. Print Vision, Ahmedabad. p.342

Singh, H. S. (2017b). Dispersion of the Asiatic lion Panthera leo persica and its survival in human-dominated landscape outside the Gir forest, Gujarat, India. Current Science, 112, 933–940. https://doi.org/10.18520/cs/v112/i05/933-940

Shaffer, J. G. & Lichtenstein, D. A. (1989). “Ethnicity and Change in the Indus Valley Cultural Tradition” In: Kenoyer, J. M. (eds.), Old Problems and New Perspectives in the Archaeology of South Asia. University of Wisconsin, Wisconsin. pp.117-126.

Staubwasser, M., Sirocko, F., Grootes, P. M. & Segl, M. (2003). Climate change at the 4.2 ka BP termination of the Indus val­ley civilization and Holocene south Asian monsoon variabil­ity. Geophysical Research Letters, 30(8), 1425. https://doi.org/10.1029/2002GL016822

Thapar, V., Thapar, R. & Ansari, Y. (2013). Exotic Aliens: The Lions & the Cheetah in India. Aleph Book Company. p.304.

Vasavada, D. T., Rana, V. J. & Ram, M. (2020). Management Plan for GIR Protected Area. Gujarat Forest Department, Gujarat, 1, p.281.

Edited By
Vishnupriya Kolipakam
Wildlife Institute of India, Dehradun, India.

*CORRESPONDENCE
Hari Shanker Singh
hss.ifs@gmail.com

CITATION
Singh, H. S. (2025). Arrival, rise, fall, and again rise of the Asiatic lion Panthera leo leo in India. Journal of Wildlife Science, 2(2), 37-45. https://doi.org/10.63033/JWLS.BHUU5534

COPYRIGHT
© 2025 Singh. This is an open-access article, immediately and freely available to read, download, and share. The information contained in this article is distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), allowing for unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited in accordance with accepted academic practice. Copyright is retained by the author(s).

PUBLISHED BY
Wildlife Institute of India, Dehradun, 248 001 INDIA

PUBLISHER'S NOTE
The Publisher, Journal of Wildlife Science or Editors cannot be held responsible for any errors or consequences arising from the use of the information contained in this article. All claims expressed in this article are solely those of the author(s) and do not necessarily represent those of their affiliated organisations or those of the publisher, the editors and the reviewers. Any product that may be evaluated or used in this article or claim made by its manufacturer is not guaranteed or endorsed by the publisher.

Anon. (1975). The Gir Lion Sanctuary Project. Government of Gu­jarat. p.131.

Barnett, R., Yamaguchi, N., Shapiro, B., Ho, S. Y. W., Barnes, I., Sa­bin, R., Werdelin, L., Cuisin, J., Larson, G. (2014). Revealing the maternal demographic history of Panthera leo using ancient DNA and a spatially explicit genealogical analysis. BMC Ecology and Evolution, 14, 70. https://doi.org/10.1186/1471-2148-14-70

Bauer, H., Packer, C., Funston, P. F., Henschel, P. & Nowell, K. (2016). Panthera leo. The IUCN Red List of Threatened Species. http://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T15951A107265605.en

Bertola, L. D., Vermaat, M., Lesilau, F., Chege, M., Tumenta, P. N., Sogbohossou, E. A., Schaap, O. D., Bauer, H., Patterson, B. D. et al. (2022). Whole genome sequencing and the application of a SNP panel reveal primary evolutionary lineages and genomic variation in the lion (Panthera leo). BMC Genomics, 23(1), 321. https://doi.org/10.1186/s12864-022-08510-y

Bertola, L. D., Tensen, L., van Hooft, P., White, P. A., Driscoll, C. A., Henschel, P., Caragiulo, A., Freedman, I. D., Sogbohossou, E. A., et al. (2015). Autosomal and mtDNA Markers Affirm the Dis­tinctiveness of Lions in West and Central Africa. PLoS ONE,11(3), e0149059. https://doi.org/10.1371/journal.pone.0137975

Clift, P. (2009). Harappan Collapse. Geoscientist, 19(9), 18–22.

Danino, M. (2016). Environmental factors in the decline of the Indus – Saraswati Civilisation. In: The Environment and Indi­an History. The C. P. Ramaswamy Aiyar Foundation, Chennai. pp.132-48.

De Manuel, M., Barnett, R., Sandoval-Velasco, M., Yamaguchi, N., Vieira, G. F., Mendoza, Z. M. L., Liu, S., Martin, M. D., Sinding, S. H. M. et al. (2020). The evolutionary history of extinct and living lions. Proceedings of the National Academy of Sciences, 117(20), 10927–10934. https://doi.org/10.1073/pnas.1919423117

Divyabhanusinh (2008). The lion of India. Black kite. Pauls Press, New Delhi.

Dutt, S., Gupta, A. K., Wünnemann, B., Yan, D. (2018). A long arid interlude in the Indian summer monsoon during ∼4,350 to 3,450 cal. yr BP contemporaneous to displacement of the Indus valley civilization. Quaternary International, 482, 83-92. https://doi.org/10.1016/j.quaint.2018.04.005

Enzel, Y., Ely. L. L., Mishra, S., Ramesh, R., Amit, R., Lazar, B., Rajag­uru, S. N., Baker, V. R. & Sandler, A. (1999). High-Resolution Holo­cene Environmental Changes in the Thar Desert, Northwestern India. Science, 284(5411), 125–28. https://doi.org/10.1126/science.284.5411.125

Fenton, L. L. (1909). The Kathiawar Lion. Journal of the Bombay Natural History Society, 19, 4-15.

Giosan, L., Clift, P. D., Macklin, M. G., Fuller, D. Q., Constantines­cu, S., Durcan, J. A., Stevens, T., Duller, G. A. T., Tabrez, A. R. et al. (2012). Fluvial Landscapes of Harappan civilization. Proceedings of the National Academy of Sciences, 109(26), E1688– E1694. https://doi.org/10.1073/pnas.1112743109

Gupta, A. K., Anderson, D. M., Pandey, D. N. & Singhvi, A. K. (2006). Adaptation and human migration, and evidence of agriculture coincident with changes in the Indian summer monsoon during the Holocene. Current Science, 90(8), 1082–1090. https://www.jstor.org/stable/24089270

Gee, E. P. (1964). The Wildlife of India. Collins, James Place Lon­don. p.192.

Gujarat Forest Department (2023a). The Project Lion @2047-A vision of Amrut Kal. Gujarat Forest Department, Gandhinagar, Gujarat.

Gujarat Forest Department (2023b). Leopard counting report. Gujarat Forest Department, Gandhinagar, Gujarat.

Gujarat Forest Department (2025). Report on 16th Lion Popula­tion Estimate. Gujarat Forest Department, Gandhinagar, Gujarat. p.36

Jhala, Y. V, Banerjee, K., Chakrabarti, S., Basu, P., Singh, K., Dave, C. & Gogoi, K. (2019). Asiatic Lion: Ecology, Economics, and Pol­itics of Conservation. Frontiers in Ecology and Evolution, 7, 312. https://doi.org/10.3389/fevo.2019.00312

Joslin, P. (1973). The Asiatic lion: a study of ecology and behavior. Ph.D. thesis. Department of Forestry and Natural Resources, Uni­versity of Edinburgh, UK, p.249.

Kazmi, R. (2021). The Last Hurrah of The Hurrianahâ Lion Pan­thera Leo Persica. Part I: A Chronological Record of Hitherto Un­known References to Lion Hunting in Present-Day Haryana and Related Reports. Journal of the Bombay Natural History Society, 118. https://doi.org/10.17087/jbnhs/2021/v118/156545

Manamendra-Arachchi, K., Pethiyagoda, R., Dissanayake, R., & Meegaskumbura, M. (2005). A second extinct big cat from the late quaternary of Sri Lanka. The Raffles Bulletin of Zoology, 12, 423–434.

O'Brien, S. J., Joslin, P., Smith III, G. L., Wolfe, R., Schaffer, N., Heath, E., Ott‐Joslin, J., Rawal, P. P., Bhattacharjee, K. K. & Martenson, J. S. (1987). Evidence for African origins of founders of the Asiatic lion species survival plan. Zoo Biology, 6(2), 99–116. https://doi.org/10.1002/zoo.1430060202

Ram, M., Sahu, A, Srivastava, N, Chaudhary, R, Jhala, L, Zala, Y. (2023a). The semi-arid ecosystem of Asiatic Lion Landscape in Saurashtra, Gujarat: Population density, biomass and conser­vation of nine wild prey species. PLoS ONE, 18(9), e0292048. https://doi.org/10.1371/journal.pone.0292048

Ram, M., Vasavada, D., Tikadar, S., Jhala, L. & Zala, Y. (2023b). Population status and distribution of endangered Asiatic lions in Gujarat, India. European Journal of Wildlife Research. 69(87). https://doi.org/10.1007/s10344-023-01720-z

Rashid M. A. & David, R. (1992). The Asiatic Lion. Department of Environment, Government of India.

Sankhala, K. (1978). Tiger! the story of the Indian tiger. Collins, London. p. 220

Singh, H. S. (2017a). The Asiatic lion-the Pride of Gujarat. Print Vision, Ahmedabad. p.342

Singh, H. S. (2017b). Dispersion of the Asiatic lion Panthera leo persica and its survival in human-dominated landscape outside the Gir forest, Gujarat, India. Current Science, 112, 933–940. https://doi.org/10.18520/cs/v112/i05/933-940

Shaffer, J. G. & Lichtenstein, D. A. (1989). “Ethnicity and Change in the Indus Valley Cultural Tradition” In: Kenoyer, J. M. (eds.), Old Problems and New Perspectives in the Archaeology of South Asia. University of Wisconsin, Wisconsin. pp.117-126.

Staubwasser, M., Sirocko, F., Grootes, P. M. & Segl, M. (2003). Climate change at the 4.2 ka BP termination of the Indus val­ley civilization and Holocene south Asian monsoon variabil­ity. Geophysical Research Letters, 30(8), 1425. https://doi.org/10.1029/2002GL016822

Thapar, V., Thapar, R. & Ansari, Y. (2013). Exotic Aliens: The Lions & the Cheetah in India. Aleph Book Company. p.304.

Vasavada, D. T., Rana, V. J. & Ram, M. (2020). Management Plan for GIR Protected Area. Gujarat Forest Department, Gujarat, 1, p.281.