Editor: Virginia Hayssen
Implications of diet composition of Asiatic lions for their conservation
Article first published online: 18 FEB 2011
© 2011 The Authors. Journal of Zoology © 2011 The Zoological Society of London
Journal of Zoology
Volume 284, Issue 1, pages 60–67, May 2011
How to Cite
Meena, V., Jhala, Y. V., Chellam, R. and Pathak, B. (2011), Implications of diet composition of Asiatic lions for their conservation. Journal of Zoology, 284: 60–67. doi: 10.1111/j.1469-7998.2010.00780.x
- Issue published online: 26 APR 2011
- Article first published online: 18 FEB 2011
- Received 16 April 2010; revised 1 November 2010; accepted 8 November 2010
- Asiatic lion;
- Gir Protected Area;
- carnivore conflict;
- livestock depredation
Livestock predation by Asiatic lions Panthera leo persica in and around Gir Protected Area (Gir PA) in western India results in conflict with people and has important implications for the conservation of this species. A 5-year study was undertaken to document diet and predation patterns based on direct observations of radio-collared lions, opportunistically located carcasses and scat analysis. Magnitude of livestock predation was assessed based on interviews of resident pastoralists in 20 settlements. Lions made one kill in every 4 days and the diet primarily consisted of large prey. Wild prey, mainly chital Axis axis, represented 80% of the lion's diet within Gir PA based on scat analysis. Within the protected area, though lions predominantly consumed wild prey in proportion to their availability, they were yet responsible for majority of livestock loss to the resident communities. The proportion of wild and domestic animals killed by lions varied between seasons: significantly more wild ungulates were killed during summer when prey were concentrated around waterholes. Domestic animals were the major prey outside the protected area. Thus, despite high proportion of wild prey in the diet, lions still considerably depended on livestock. Our study defines focal areas of lion–human conflict and suggests better husbandry practices.
Population decline, crisis management, stabilization, precarious recovery and sustained recovery have been described as five stages of species restoration (Linklater, 2003). Currently, carnivore conservation involves a sixth stage beyond population recovery, that of managing successful recoveries of target species that exceed the carrying capacity of protected areas (Hayward, O'Brien & Kerley, 2007a). During this stage, space and conflict mitigation become the principal conservation concerns (Macdonald & Sillero-Zubiri, 2002; Inskip & Zimmermann, 2009). Among these issues, livestock predation is the most challenging (Macdonald & Sillero-Zubiri, 2002). To assess the magnitude of such conflict, knowledge of predator diet is crucial (Hayward & Hayward, 2006), especially in countries like India, where people and wildlife live in close proximity to each other and livestock predation causes significant economic loss.
Predation on livestock by large carnivores is variable (Mukherjee & Mishra, 2001; Biswas & Sankar, 2002; Bagchi, Goyal & Sankar, 2003; Andheria, Karanth & Kumar, 2007) and governed by availability and vulnerability of livestock and wild ungulates. In areas of substantial wild ungulate densities, tigers consumed smaller proportions of livestock (Biswas & Sankar, 2002; Andheria et al., 2007) while in other areas, in spite of high prey abundance, they consumed considerable numbers of livestock that were readily available within the protected area (Mukherjee & Mishra, 2001; Bagchi et al., 2003). In wild prey-deficient habitats, while leopards switched to a diet of domestic prey in some areas, tigers preferentially killed smaller wild prey and avoided killing livestock in spite of their availability within the park (Edgaonkar & Chellam, 2002; Reddy, Srinivasulu & Rao, 2004). Availability of livestock in a protected area thus does not necessarily represent the magnitude of conflict between carnivores and local communities. Instead, examination of predator diet and expression as proportion of livestock and wild prey consumed would be a better indicator and also help to overcome the difficulty of quantifying ‘availability’ of guarded domestic prey.
Assessment of diet and prey preference of Asiatic lion Panthera leo persica is important for conservation and management in this scenario of increasing lion population, change in land-use, increasing human population and the ensuing conflict (Pathak et al., 2002; Vijayan & Pati, 2002; Meena, 2010). We undertook a study to estimate (1) Lion diet and predation pattern; (2) Livestock losses to predation to understand the magnitude of human–lion conflict.
Materials and methods
Gir Wildlife Sanctuary (1153.4 km2) and National Park (258.2 km2) constituting the Gir Protected Area (Gir PA) is located in the southern part of the Kathiawar peninsula, in the state of Gujarat in western India (Fig. 1), at 21°20′ and 20°57′N latitude and 70°27′–71°13′E longitude. Gir has a semi-arid climate with average temperatures ranging from 10 to 43 °C, with an average rainfall of 900 mm and with three distinct seasons, hot and dry summer (March to mid-June), monsoon (mid-June to mid-October) and cool and dry winter (late October to February). Vegetation is tropical dry deciduous forest interspersed with tropical thorn forest (Champion & Seth, 1968). Gir PA is divided into three management units, namely Sanctuary West (SW), National Park (NP) and Sanctuary East (SE) that vary with respect to rainfall, topography, vegetation, management regimes and anthropogenic pressures (Khan et al., 1996; Singh & Kamboj, 1996). While NP is inviolate, around 12 334 livestock are resident in forest settlements and nesses (hamlets of local pastoral community, the Maldharis, within the protected area) and permitted to graze in SW and SE (Fig. 1, Pathak et al., 2002). Data on prey carcass and scats were collected in SW, SE, NP and areas falling c. 5 km outside protected area boundary (hereafter referred to as peripheral areas). Approximately, 94 582 livestock (chiefly buffalo and cattle) are present in the 97 peripheral villages falling within this zone. Ness survey and monitoring of feeding habits of individual lions was carried out within an intensive study area of 1075 km2 covering SW and NP (Fig. 1).
The only surviving free-ranging population of Asiatic lion exists as a single population in and around the Gir PA. Lions underwent a population bottleneck more than 100 years ago (O'Brien et al., 1987) and subsequently passed through the five stages of conservation mentioned (Linklater, 2003). The Gir Lion Project (1972) to revive the population of Asiatic lions, implemented stringent conservation measures including partial removal of people and livestock out of the protected area. The project helped to check the increasing animosity among livestock owners towards lions as a consequence of enormous losses to predation and inadequate compensation (Joslin, 1973). During this period lions lost their livestock kills to hide collectors and also succumbed to occasional carcass poisoning (Joslin, 1973). As a result of successful management there has been an increase in the lion population from <50 (Dalvi, 1969) at the turn of the last century, to about 411 in 2010. An estimated 114 lions occur in Girnar Sanctuary, Mitiyala Sanctuary, Savarkundla, Liliya and adjoining areas and constitute the ‘satellite lion population’ (Meena, 2010).
Apart from lions, other large carnivores in Gir include leopard Panthera pardus and striped hyena Hyaena hyaena. Wild prey comprises of chital Axis axis, sambar Rusa unicolor, nilgai Boselaphus tragocamelus, chousingha Tetracerus quadricornis, chinkara Gazella bennetti, wild pig Sus scrofa, porcupine Hystrix indica, common langur Semnopithecus entellus, rufous tailed hare Lepus nigricollis ruficaudata and peafowl Pavo cristatus (Singh & Kamboj, 1996). Prey are resident throughout the year with minimal seasonal variation (Khan et al., 1996). Wild ungulate density (±se) is estimated at 48.3 (±6.1) individuals km−2 (Table 2; Dave, 2008).
|Year||Prey type||Prey density (no. per km2)||Frequency of occurrence in lion diet based on scat analysis (%)||Source|
|Livestock (resident)||21.5 (SE) 9.8 (SW)||25.1|
|2002–2006||Overall||48.3||Dave (2008); Meena (2008)|
|Livestock (resident)||6.2 (SW)||19.9|
Determination of diet composition and predation pattern
Data collection was carried out from April 2002 to December 2006, except for continuous day–night observation on radio-collared lions that was carried out only in 2006. Data collection involved a combination of observations on six radio-collared lions, kill remains from opportunistic searches and scat analysis. Questionnaire surveys were administered to collect data on livestock losses from settlements within the protected area. Diet variation was assessed for the differentially managed zones.
Odour, prey alarm calls, presence of crows and lion signs (tracks and drag marks) were used to locate prey carcasses (hereafter referred to as kills). The distinction between lion and leopard kills was based on evidence around the kill such as pugmarks and predator hair, mode of feeding and state of kill remains (Chellam, 1993). Asiatic lions are social predators consisting of female prides and male coalitions that hunt and feed independently (Meena, 2009). Lions typically rip apart, scatter carcass remains when feeding together and eventually completely consume the prey, leaving nothing edible behind. Leopards on the other hand, start feeding from the rump, hide the rumen sac and cache kills (Chellam, 1993).
Frequency of occurrence of a prey species was calculated as the number of times a specific prey item occurred and was expressed as percentage of all prey occurrences. Seasonal diet variation as well as differences in diet between different geographical areas of the protected area was tested using χ2 analysis (Zar, 1999).
Lion scats were collected mainly along roads and forest tracks. Lion scats were clearly distinguishable from leopard scats based on their much larger size. Nevertheless, carnivore signs associated with scats were additionally recorded. All scats were stored in tagged polythene bags and later washed using a sieve to separate undigested prey remains such as hair, bone fragments, hooves, feathers, quills and claws. All remains were oven-dried for further examination.
For a reliable estimate of lion's diet, standard prescribed protocols – examination of a minimum of at least 20 prey hairs per scat and minimum 30 scats – were adopted (Mukherjee, Goyal & Chellam, 1994; Jethva & Jhala, 2003). Microscopic slides of randomly picked hair from a sample were washed in xylene and examined under a light microscope. Prey were identified by comparing medullary characteristics of prey hair with known standard reference hair (Karanth & Sunquist, 1995).
Frequency of prey obtained from analysis of scats was subjected to re-sampling to obtain confidence limits on the mean percentage of prey in the scats (Reynolds & Aebischer, 1991). This involved iterating sub-samples of the same size 10 000 times using bootstrapping in the computer programme simstat (Peladeau, 1995).
Representation of prey intake as per cent frequency of occurrence of the seven major prey species based on scat data can be misleading due to variation in relative contribution of various prey species that vary with varying body size. A bias occurs due to the surface area to volume relationship, whereby smaller mammalian prey produce relatively more scats per kilogram of meat consumed than do larger prey, leading to overestimation of small prey in predator diets. This bias can be overcome by expressing prey intake as relative biomass and relative number of prey taken (Floyd, Mech, & Jordan, 1978). For estimating relative importance of the prey species, a correction factor developed for cougar Felis concolor (Ackerman, Lindzey & Hernker, 1984) was applied by assuming that lion digestive physiology is similar to that of the cougar's. The regression equation used is y=1.980+0.035x, where y is the biomass of prey consumed (kg) to produce a single field collectable scat and x is the average body weight of the prey species (kg). This relation was used to convert frequency of prey occurrence in scats into relative biomass and number of prey consumed.
Observations of radio-collared lions
Direct observations were made on 10 feeding events of six radio-collared lions, three males and three females, to supplement opportunistic recordings of kills. Five sessions of continuous day–night observations ranging from 5–10 days, totaling 38 days, on three radio-collared males belonging to three different coalitions was carried out.
Survey of livestock holding and predation
A survey was conducted in all the 20 resident nesses and settlements within the intensive study area to obtain information on annual (2003–2004) livestock loss to lion predation. A total of 148 families were interviewed that included 1408 resident forest dwellers to collect information on number of families, number of individuals per family and livestock-holding in each household. Information on livestock mortality was classified as loss due to predation and loss due to other natural causes and percentage loss due to predation was calculated.
Data on 1215 lion attacks on livestock from January to December 2006 was obtained from Gujarat Forest Department to examine the time of attack.
Selectivity indices, such as Jacobs index with values ranging from +1 (maximum preference) to −1 (maximum avoidance) indicate diet preference taking into account both proportion of kills and prey availability (Jacobs, 1974). Hayward & Kerley (2005) derived Jacobs index scores (D) for major lion prey species from Jacobs index preference equation (Jacobs, 1974):
where r is the proportion of kills and p is the proportional abundance of each prey species. Hayward et al. (2007b) further derived an equation for predicting the number of kills of a particular species based on the above equation.
where Ri is the predicted number of kills of species i when a total of ∑K kills are observed, Di represents the Jacobs index value of species i and pi represents the proportional abundance of prey species i at a site.
Prey preference was modelled for 2002–2006 (present study) by obtaining Jacobs index scores (D) for four major lion prey species of Gir, namely, chital, sambar, nilgai and wild pig from Hayward & Kerley (2005) and deriving predicted number of kills of each of these species based on proportional abundance of each prey species (Dave, 2008) and proportion of kills observed (Table 3). The accuracy of the model prediction was tested using the log-likelihood goodness of fit (G) test (Zar, 1999).
|Species||Jacobs index (D)||Proportional prey abundance (p)||Kills of 2002–2006|
|Observed (k)||Predicted (R)|
|Chital Axis axis||−0.81||0.901||72||64|
|Sambar Rusa unicolor||−0.16||0.059||26||5|
|Nilgai Boselaphus tragocamelus||−1||0.025||6||0|
|Wild Pig Sus scrofa||−0.39||0.016||15||1|
Of 258 kills, livestock constituted 53% and wild prey 47%. Cattle were 31% of the total, chital 28%, buffalo 16%, sambar 10%, nilgai 3%, wild pig 6%, goat 3%, camel 2%, peafowl and chousinga 1%. Proportion of wild kills in summer was 67% (n=100), 35% (n=68) for monsoon and 38% (n=90) for winter.
Predation on wild and domestic prey varied between peripheral areas, Gir West and NP (χ2=12.3, d.f.=2, P<0.001). Livestock was part of the diet in all areas of the protected area: 38% of kills in NP (n=46), 50% of kills in Gir West (n=170) and 76% of kills in peripheral areas (n=42). Consumption of wild and domestic prey varied between seasons (χ2=22.3, d.f.=2, P<0.0001) with a greater proportion of wild prey killed during summer months (Fig. 2).
Three hundred and ten lion scats were analysed and 12 prey species were identified. Two hundred and ninety-five (95.2%) scats had a single prey item while 15 (4.8%) scats had two prey species. Frequency of prey occurrence were: chital 32%, sambar 26%, wild pig 10%, buffalo 11%, nilgai 9%, cattle 8% langur 2% and minor prey 2% including peafowl, porcupine, an unidentified bird species and camel (Table 1). Remains of claws of felid cubs were found in two scats but we were unclear as to whether they belonged to lion or leopard claws. Diet did not differ among management zones nor among seasons. Frequency of occurrence of prey in scats revealed that livestock occurred in only 20% of the scats and wild prey occurred in 80% of the scats (Table 1). Chital and sambar contributed up to 58% of the lion's diet (Table 1).
|Species||Weight (kg) (Schaller, 1967)||Frequency of occurrence (%)||Collectable scats per kill||Bootstrapped CI (95%)||Relative biomass consumed (%)||Relative number of individuals consumed (%)|
Observations on radio-collared lions
Mean (±sd) feeding interval of 0.26 (0.055) kills per day translated to one kill every 3.9 that is, 4 days. Eighty per cent of the prey consumed were adults constituted by chital (three), sambar (two), cattle (two) and buffalo (three). Fifty per cent of kills occurred between 16:30 and 20:00 h.
Survey of livestock holding and predation
Fifty one per cent of 3896 buffaloes and 43% of 847 cattle were adults (Fig. 4). Of the total annual livestock mortality, 60% (n=361) was due to lion predation and the rest due to disease and old age. Adults formed 49% (n=102) of buffalos and 69% (n=89) of cattle lost due to predation. Most of lion attacks (34%) on livestock occurred between 16:00 and 20:00 h (Fig. 3).
The model accurately predicted the observed number of kills for the period 2002–2006 (G=11.4, d.f.=2, P=0.05) (Table 3).
The success of Gir Lion Project was reflected in recovery of native vegetation as well as increase in wild ungulate and lion populations (Table 2; Khan et al., 1996). This success contributed to a dramatic change in the lion's diet (Table 2; Chellam, 1993). In the past, livestock formed 75% of the lion's diet (Joslin, 1973). However, during the subsequent period 52% (in early 1980s) and 75% (in the late 1980s) of prey consumed were wild prey (Sinha, 1987; Chellam, 1993). Similar ecosystem revival and management interventions need to now extend beyond protected area boundary to ensure conservation of lions in the long term in the entire landscape. We therefore discuss the findings of our study based on a framework of a three-pronged management strategy to ease conflict: first, identification of focal regions for intervention having greater incidences of livestock predation; second, improving husbandry practices based on a better understanding of both lion predation patterns as well as traditional methods of vigilance; third, offering appropriate compensation for economic losses incurred due to lion and leopard predation.
Identification of conflict areas
Owing to the greater availability of livestock, particularly cattle, in the peripheral areas as compared with resident livestock, a clear-cut difference in lion diet was evident within and outside protected area. Within the protected area also, including the NP which is located within the Gir PA (Fig. 1), livestock formed a significant part of lion's diet. Livestock constituted 47% of lion diet within the Gir PA while in the peripheral areas, livestock constituted 76% of the 42 kills. Compensation claim records of the Forest department also indicated that average livestock loss to predation per month within protected area to be 45 and outside protected area to be 89 (Pathak et al., 2002). Livestock remains were found in 21% of 29 kills collected from NP, 43% of 117 kills of SW and 69% of 32 kills outside protected area (Chellam, 1993).
Livestock owners residing within 5 km of Gir PA do not have clear-cut grazing rights and therefore benefit less from proximity to the forest. Yet, more livestock predation occurs outside the protected area because of greater availability of livestock, low density of wild prey (mostly nilgai), and increased lion movement (Soni, 2000; Pathak et al., 2002; Meena, 2010). Thus, focal areas of interventions have to be outside the protected area.
Abundance, size and temporal and spatial distribution of prey influence hunting strategy, activity and daily movement of lions (Schaller, 1972; Eloff, 1973; Stander, 1991; Patterson et al., 2004). Gir has high biomass of resident wild prey available throughout the year in addition to availability of relatively more vulnerable domestic livestock prey base. Felids require large prey and African lions Panthera leo leo preferentially prey upon species of an average weight of 350 kg, range 190–550 kg (Hayward & Kerley, 2005). Our study also indicates greater consumption of large-sized prey in adult age class (Fig. 2). Although, incidental observations of kills tend to be biased towards large bodied prey because of easier detectibility, our kill data represented by 62% large bodied wild prey are yet comparable to findings from scat analysis. Monitoring lions with the help of radio-telemetry confirmed that 80% of kills (n=10) were of adult prey. Overall, in terms of relative number of individuals consumed, domestic prey occurred in low proportions (20%) yet in terms of biomass contribution, they accounted for 36% (Table 1).
In the wild, lions have to hunt to meet their daily requirement of 5–7 kg (Schaller, 1972). In captivity, Asiatic lions (average body mass 100 kg) consume 6% of their total body mass as buffalo meat in a day (Mukherjee & Goyal, 2004) while in the wild, they consume 7–10% of their body weight (Mukherjee & Goyal, 2004). To meet this requirement, a single lion would have to consume 8–12 kg of meat in a day that in turn translates to 240–360 kg per month. Taking into account their daily requirement, prey body weight (Table 1) and prey preference, a single lion would have to kill two cattle or one buffalo per month. Official records indicate 90 livestock kills occur each month that in turn implies that a maximum of 45 lions (15% of the population) are totally dependent on livestock predation.
In places where lions depend on livestock, they resort to nocturnal predation (Schaller, 1972; Van Orsdol, 1984; Patterson et al., 2004). In Gir, because the livestock were well protected within stone fences and corrals at night, predation occurred mostly between 16:00 and 18:00 h, when livestock were brought back from their foraging grounds (Fig. 3).
Among wild-prey, chital was the most commonly killed species (Table 1). Proportion of wild ungulate kills was greater in summer (67 of 100 kills) probably due to greater hunting success around localized water sources. An increase in adult stag kills, particularly chital, occurred in winter during rutting season (Fig. 2). Wild prey predation occurred between 16:30 and 20:00 h. Lions made one kill every 4 days and also scavenged on dead, sometimes even decaying prey and snatched kills from leopards (n=13). Some individual lions, particularly older males depended largely on livestock predation or on scavenging and appropriating kills from lionesses or leopards (V. Meena, pers. obs.). By constant vigilant monitoring, such individual lions predating largely on livestock, could be selectively captured as suggested by Hemson (2003). The prey preference model accurately predicted predation patterns during the period 2002–2006 for Asiatic lions. Although livestock consumption is not included, the model accurately predicts consumption of wild prey that corresponds to observed changes in diet. In Gir, wild prey is consumed in proportion to availability without specific preference. Hayward et al. (2007b) have further extended these models to predict carrying capacity of large predators in conservation areas and these may be applied for predicting carrying capacity in and around Gir PA in the future.
Historically, while the tolerance among livestock owners has fluctuated with time, lions have always preyed on livestock (Joslin, 1973). Thus, conservation measures should address the lion's dependency on livestock. Improving husbandry practices may reduce losses at least at an individual herd-level. Based on observed predation patterns following preventive measures can be implemented such as increased vigilance during evening hours, restricted grazing or stall feeding and decrease in livestock holding by maintaining fewer but more productive breeds.
For livestock owners, low monetary investments and high profit margins obtained from animal husbandry appears to offset overall loss due to predation. Overall, predation accounted for only 4% of the total livestock population lost annually. Our survey revealed that on an average (±sd) each family owned 13 (11) buffalos and 2.5 (3) cattle. Of the total, 33% of buffaloes and 43% of cattle were productive adult females in that year. Only four buffaloes and one cow per family were productive. Most of the livestock kills recorded were of adults with high market value – buffalo 10 000–30 000 rupees; cow 2000–8000 rupees. Livestock predation thus causes considerable economic loss contributing to 60% of annual livestock mortality. Therefore, losses incurred due to predation are compensated by the Gujarat Forest Department at rates calculated and revised periodically to reduce disparity between market price of animal lost and compensation offered. In spite of this the instant financial incentive provided by monetary compensation to help reduce impact of loss due to predation, the cultural implications and emotional costs cannot be accounted for (Macdonald & Sillero-Zubiri, 2002). Therefore, people's tolerance of livestock losses cannot be sustained by monetary compensation mechanisms alone.
Human–carnivore conflict, particularly due to livestock predation is a global issue with no permanent solution. Based on our study, much of this conflict is outside Gir PA, in private lands where neither livestock nor the owners can be moved or resettled. Improved husbandry practices based on ecological information on lion's diet such as prey preference and time of attack in combination with the suggested livestock management practices and monetary compensation would be required for the continued positive attitude of local communities and long-term lion conservation.
This project was funded by the Wildlife Institute of India. We are grateful to the Ministry of Environment and Forest, Government of India and Chief Wildlife Warden of Gujarat for permissions and facilitation. We would like to thank K. Bannerjee and field assistants, Biku, Taju, Ismail & Guga for helping in field data collection. We thank Vinod Thakur for guiding and helping with laboratory work. We are grateful to Shomita Mukherjee Manoj Nair, Mathew Hayward and unknown referee for critical comments and help with improving the paper.
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