understanding tiger ecology in the tropical dry forest of

1

Understanding Tiger Ecology in the Tropical DryForest of Panna Tiger Reserve

Study Period: 1996-2002

Dr. Raghunandan Singh Chundawat, Principal InvestigatorMr. Neel Gogate, Researcher (1996-1999)

Dr. Pradip K. Malik, Co-Investigator

Global Tiger Patrol, UKNational Fish and Wildlife Foundation

Wildlife Institute of IndiaWildlife Conservation Society

First D

raft: fo

r inter

nal use

and

not for

circulat

ion

Dated: 28th September 2005

2

AcknowledgementsI am very grateful to Shri P.K. Mishra, and Shri A. P. Dwivedi, PCCF and CWLW, Govt. Madhya Pradesh for provid-

ing support and permission to carry out the fieldwork in Panna National Park. I also thank Shri H. S. Pabla and ShriSuhas Kumar and the Staff of the Chief Wildlife Warden office for their help.

The fieldwork of the research project could not have achieved what it has without the help and support of ShriP. K. Chowdhary and Shri Sanjay Mukharia, Field Directors of Panna National Park and their staff.

The Research Project was funded by the Wildlife Institute of India, Dehra Dun initially and later by Global TigerPatrol, UK, and the National Fish and Wildlife Foundation USA and supported by the Wildlife Conservation Society,USA. I thank them for providing me the opportunity to understand the ecology of tigers in a Dry Forest. I also thank theDirector, Wildlife Institute of India, Dehra Dun for allowing me to take leave and work on the project.

The project was supported by several other organisations and individuals in many ways. I am grateful to all theconcerned individuals of these organisations. The Wildlife Conservation Society and U. S. Fish and Wildlife Serviceprovided helped procuring equipment. Ranthambore Foundation provided a four-wheel drive vehicle for the project. TheCentre for Wildlife Studies, Bangalore provided logistic support and helped the project in all possible ways, includingcollaborating in conducting the line transect surveys in March 2001. I particularly thank Dr. Ullas Karanth and ShriSamba Kumar and the trained volunteers from the Centre for Wildlife Studies.

In Panna, the project work had benefited from the help of many individuals; I wish to thank Shri Dave Ferguson ofthe U. S. Fish and Wildlife Service for his support and help. I also thank the entire staff of the Panna National Park fortheir co-operation and help to the project, and especially the elephant mahouts, without whose support very littlecould have been achieved.

Several individuals also showed a keen interest in the research project and supported it whenever a need ar-rived. For their help and encouragement, I thank Shri S.C. Sharma, Additional Secretary (WL) MOEF, Shri P.K. Sen,Director, Project Tiger, Shri Suhas Kumar, CF, Office of Chief Wildlife Warden, Shri Dharmendra Shukla, Shri P. K. Chowdhary,previous Field Director, Panna National Park and Dr. A.J.T. Johnsingh at the Wildlife Institute of India. I am highlythankful to Shri Koustubh Sharma and his field staff for his enormous help in conducting some of the fields exercisesand data analysis.

Dr. P.K. Malik and his veterinary colleagues were of course invaluable to the project and I thank them deeply: theyshared their expertise and enabled us to successfully radio-collar the animals in Panna. Much of the work discussed inthis report owes thanks to Dr. Peshin and Dr. A. B. Srivastava.

In particular I would also like to thank Ms. Amanda Bright and Mr. Toby Sinclair of Global Tiger Patrol, Ms. BelindaWright of WPSI, Shri Valmik Thapar, Shri P. K. Sen, Dr. John Seidensticker, Ms. Tracy Walmer of National Fish and WildlifeFoundation, Shri Sanjay Gubbi of the Centre For Wildlife Studies, Ms. Priya Ghosh and Ms. Alice Pandya from theAmerican Embassy, Ms. Uttara Mendiratta, Faiyaz Khudsar, Vijay Panday; Shyamendra Singh and Bhavana, Shri VishalSingh, Mr. Julian Mathews of Discovery Initiatives their personal efforts provided critical help that made this workpossible.

The fieldwork would not have been possible without the help of the research staff, who worked tirelessly andlaboured every day to help me collect the information for the project. I feel very fortunate to have had such a wonderfulteam. The success of the research project comes mainly from the hard work put in by them. I thank all the fieldassistants namely Shri Kalyan Singh Yadav, Shri Bansh Gopal Yadav, Shri Mahadev Yadav, Shri Suresh Singh Yadavand Shri Hargovind for their hard work and sincere dedication for the research project. I also thank Shri Sataru, ShriMadan Singh Yadav and Shri Dayaram Yadav, project staff members, who worked with the same sincerity and dedica-tion but are not with the project now. Shri Dhan Bahadur was also a vital part of the team. I also sincerely thank ShriR.T. Sanago, Range Officer on leave, who worked for his Ph.D. on the ecology of sambar as part of the project.

At the end I wish to thank my wife, Joanna Van Gruisen, who accompanied me in the field and whose vastexperience and understanding of Indian jungle life was of tremendous help to me everyday. She is my most rigorousreviewer and I am grateful to her also for the hours she spent editing the draft of this report.

3

Understanding Tiger Ecology in the Tropical Dry Forest of PannaTiger ReserveIntroduction

Among all wild animals, the tiger hasreceived a considerable amount of attention theworld over. Despite such favourable notice, tigerpopulations are declining at a faster rate thanat any time in their history. Many local isolatedpopulations are being pushed towards extinction.The recent loss of tigers from Sariska is one suchexample that highlights the extreme vulnerabilityof tiger populations in the Indian sub-continent.The decline in tiger numbers is mainly due toincreasing human pressure on tiger habitatsthroughout its range; mounting demand onnatural resources is destroying the tigers’habitat and increased man-wildlife conflictscreate an environment unsuitable for the survivalof a large predator like tiger. In these habitatsused by humans, the tigers’ survival is precariousand in most places, it is on the verge of localextinction. The situation is worse outside theprotected areas.

Since enacting the Indian Wildlife(Protection) Act, 1972 and the inception of“Project Tiger” in April 1973 in India, considerableeffort has been made towards the conservationof tigers in India (Panwar, 1982, 1987). Therepeated occurrence of tiger crises indicatesthat not enough has been done and the effortsmay have been imperfectly directed; certainlymuch more needs to be done to save wild tigersin India. A network of protected areas, as TigerReserves, supposedly safeguards extensivewilderness especially for the tiger. Yet, the tigerpopulation within these protected areas can alsobe affected by several ecological factors inaddition to the existing human induced bioticpressures and developmental activities such aslarge dams. Like all large carnivores, the tigerrequires adequate space, substantial prey base,cover and protection from human causedmortality (Schaller, 1972; Gittleman and Harvey,1982; Mills, 1991; Caro and Durrant, 1995). Tiger populations, in most of the protectedareas in India, face the following severalconservation problems (Panwar, 1982, 1987;Wemmer et al, 1987, Karanth, 1991; Johnsinghet al, 1991):

a) Inadequate size of protected areas.b) Fragmentation and loss of habitat, as well

as an increasing rate of habitatdegradation resulting in a decrease inhabitat quality.

c) Inadequate prey base.d) Mounting demands for forest based

resources, thereby increasing man-wildlifeconflicts.

e) More recently, poaching for commercialuse.

As a result, the tiger only survives in smalland isolated populations (Johnsingh et al, 1991).Such small populations are vulnerable to normallyoperating demographic and environmentalstochasticity (Caughley, 1994). This vulnerability,in addition to the factors mentioned above, makesuch populations very unstable (Soule 1986); aslightest perturbation in the ecosystem canhave a pronounced effect on the stability and inturn on the viability of these populations.Therefore, the overall survival of tigers will dependupon the stability and viability of localpopulations.

Problem StatementTo maintain a demographically viable

population of tigers, the size of the protectedarea is the critical issue for management(Woodroffe and Ginsberg, 2000). The rate ofextinction for any larger mammal species is rapid,and is more pronounced in the case of a largecarnivore population in small and isolatedprotected areas. There are no defined methodsto determine what should be the minimum sizeof a protected area (May, 1975; Terbough, 1976;Simberloff and Abele, 1976). Yet managementof a protected area should strive to conservethe species for which it has been created (Smith,1984). It would, therefore, be appropriate toconsider tiger as an indicator species to addressthe issue of size of protected areas and otherecological problems that exist in its habitat.

In India, most of the protected areas that

4

have tigers, are small and isolated units, eitherdue to their physical fragmentation or becauseof managerial differentials (Panwar, 1987;Johnsingh et al, 1991). Approximately 80% ofthe protected areas in India are less than 400km2 in size (Rodgers and Panwar, 1988). In sucha situation, it is important for management todetermine the potential of PAs in conserving tigerpopulations. In other words, based on theanimal’s ecological requirements, it is importantto consider approximately how many tigers a PAcan support. Rather than simply plotting thetiger population growth chart, such an approachwould be more meaningful for the conservationof this endangered species. A field-orientedapproach of this type would eventually give arealistic projection of the requirements neededto maintain demographically viable populations.In effect, it would improve our scientificunderstanding for managing protected areas forthe conservation of tiger populations.

The pertinent question that Managementneeds to ask is: ‘are the present protected areascapable of sustaining a prey base, which cansupport a demographically viable tigerpopulation?’

How does Management go aboutanswering this question?a. Like most other carnivores, tigersspecialize in their feeding habits; their predatorystrategies and have evolved through a rigorous

natural selection process to maximize theirenergy intake (Sunquist and Sunquist, 1989).Therefore, the ecology, social organization andmovement pattern of tigers are likely to begoverned by several ecological factors such asavailability of prey, its temporal and spatialdistribution, its size and anti-predatorybehaviour. It, therefore, becomes necessary tocollect information on tiger feeding ecology,ranging and movement pattern to find answersto the question asked earlier.b. Animals occupy an array of habitats, andtheir demography varies with the differenthabitats type they occupy. The reproductivesuccess of a population largely depends on theproportion of suitable habitat available forbreeding. In “source” habitats, where surplusindividuals are produced, any loss or alterationto these habitats due to natural oranthropogenic reasons will have a drastic effecton the stability and survival of the population(Pulliam, 1988; Pulliam and Danielson, 1991).Therefore management needs to know - how arethese “source” habitats utilized by tigers? Whatare the characteristics of such habitats? Howdo we identify and manage these habitats tomaximize the reproductive success of tigers?These are a few of the important managementquestions that need immediate answers andthat will benefit management in achieving its goalof tiger conservation.

c.A n o t h e rimportant issuethat needs to beaddressed is thatof the dispersalpatterns of juveniletigers and theirsurvival. Dispersalnot only regulatesthe population butalso maximizesmating success andplays a major role ingenetic flow andsocial organization(Dobson, 1982;Smith, 1984).Populations in a“source habitat”

5

(resource rich) produce excess individuals, whichthen have to disperse from their natal areas intoneighbouring “sinks” (sub-optimal habitat). Themanagement of dispersing individuals in thesesinks is extremely important and small PAs as“source” habitat need to be linked to theneighbouring “sinks”, so that tiger populationscan be conserved as a meta-population (Pulliam,1988). A study of such dispersal dynamics wouldprovide scientific guidelines for the managementof meta-populations, which in India is longoverdue. The issues discussed above are consideredpertinent for this study and it attempts toanswer following questions:1. What is the food requirement for tiger?2. What is the minimum size of protected

area required to sustain a demographicallyviable population of tigers?

3. Does tiger prey regulate its social system,ranging pattern and habitat requirement?

4. What are the “source” habitats and howare they utilized?

5. What happens to the individuals thatdisperse?

6. How do tigers react to human interferencein “sink” habitats?

7. What would be the best conservationstrategy for managing large carnivorepopulations in small isolated protected

areas?

Need for the studyOver 60% of the world’s tiger population

survives within the Indian subcontinent. The tigeroccupies a wide range of habitats in India butvery few protected area are large enough toprotect a large viable population (Rodgers andPanwar, 1988). Although, in the past a fewstudies have been conducted on tiger (Schaller,1967, Seidensticker, 1976, Sankhala 1977,McDougal, 1977, Panwar, 1979, Sunquist, 1981,

Tamang, 1982, Smith, 1984), most of them werein its optimal habitat. Thus most of theknowledge available on tiger ecology is from moistand productive habitats.

On the other hand the tiger census figuresof 1993 report that about 50% of the Indiantiger population is living outside the ProtectedAreas, which are sub-optimal habitats for tigers.The average tiger population in these PAs is lessthan10 tigers. At present about 45% of theremaining tiger habitat in the Indiansubcontinent, is in Tropical Dry Forests(Wikramanayake et al 1999). Extremetemperatures, seasonality, low productivity andhigh human pressure characterize thesehabitats. The tiger has suffered mostly in thesehabitats. About 35% and 70% of the tigerpopulation has become locally extinct from DryDeciduous Forests and Semi-arid Foresthabitats respectively. Such large-scale localextinctions of tiger population in Dry Foresthabitats highlight the vulnerability of tigerpopulations in its largest habitat in India.Therefore, this study was conducted in a DRYFOREST HABITAT, which represents a typicalscenario, where tigers experience all the problemsdiscussed above. The Panna National Park/TigerReserve of Madhya Pradesh was selected for thestudy.

OBJECTIVES OF THE STUDY were to:* evaluate habitat suitability for tigers and

assess the availability and distribution ofits major prey;

* estimate and study the home ranges andmovement patterns, land tenure patterns,socialorganization and habitat use bytigers;

* determine food habits and foodpreferences of tigers;

* develop a model to suggest managementmeasures to protect tiger populations inDry Tropical Forests.

6

Panna National Park, in the central Indianhighlands, represents one of the important tigerhabitats of the Tropical Dry Forest (Chawdhry1997; Chundawat et.al 1997; Gogate andChundawat 1997; Pabla, 1984). Panna NationalPark along with the surrounding territorial forestareas is the only large tiger habitat remaining innorth Madhya Pradesh (M.P.). These forests werethe private game reserve of the erstwhile princelystates of Bijawar, Chattarpur and Panna. Apartfrom the conservation values, these forests arealso home to several rock paintings and templesdating back 2000 to 4000 years (Chaudhary,1996). The Panna National Park attained the legalstatus of Protected Forest in 1948. Part ofthese forests was declared National Park in 1981;this included a major part of the existing GangauWildlife Sanctuary. The M.P. State Government,issued a notification dated 17-10-1981, todeclare this area as National Park; this waspublished in the State Gazette of 1.1.1982 (M.P.Gazette, 1985); the final notification undersection 34(4)(A) of the Act is still awaited.

Subsequently, in 1994, Panna National Park wasidentified as an important tiger habitat anddeclared a Project Tiger area. The total area ofthe park is 542.66 km2. For administrativepurposes, the park is divided into fourmanagement units ? ranges, viz: Panna, Hinouta,Madla and Chandranagar. The Park headquartersare situated at Panna town. Panna National Park is situated in thenorthern extension of the Vindhyan rangesbordering UP in the north (Chaudhary 1996). Itspreads over two of M.P.’s districts: Panna andChhattarpur (Figure 1). Biogeographically andfloristically, this area forms part of the Indo-Malayan Region, whereas zoo-geographically, itis a part of the Oriental Region (Mackinon,199XX). It lies in Zone 6 E, i.e., ‘Deccan Peninsula- Central Highlands’ (Panwar and Rogers, 1988).The location of the National Park is alsoimportant because it is situated at a point wherethe continuity of the forest belt gives way tothe extensive Gangetic Plains (Pabla, 1984). The topography of the area is characterizedby step tablelands with two distinct plateaux

The Study Area: Panna Tiger Reserve

7

above the Ken river valley. Altitude varies from220m at the river bank to 540m. The fall betweenupper and middle plateau and middle plateau andthe Ken river valley is steep, forming 10-80m highescarpments. This distinctive landscape givesrise to many gorges, cliffs, and overhangs. A largenumber of caves and rock shelters, springs andpatches of thick forest cover are also spreadover the area, creating unique microhabitats fordifferent animals and birds.

The Vindhyan strata in this region areknown for their poor water retaining capacity dueto the large number of fractures and faults. Asa result, most of the surface water percolatesdown very fast, leaving streams, nallahs andpools dry soon after the monsoon. Streams inthe National Park are seasonal; the Ken river isthe only perennial river. It flows for about 55 kmthrough the park, before joining the Yamuna riverin Banda district of U.P.

The climate of the area can be describedas typically dry tropical with three distinctseasons. The summer begins in March and lastsuntil the first shower of the monsoon in June.May is the hottest month of the year andtemperatures often rise as high as 48° C. Theannual rainfall averages about 1100 mm andmost of it ? over 70% ? falls during July andAugust only. Consequently, a long dry spellfollows the monsoon. Winter is short, buttemperatures can regularly go below 5°C andfrost is a common feature in open habitat areas.

According to Champion and Seth(1968), this area is classified as Northern DryDeciduous Teak Forest (5 B C/2). The dominatingvegetation type is Miscellaneous typeinterspersed with grassland areas. Other majorforest types are riverine, open grasslands, openwoodlands with short grasses, closed woodlandswith short and tall grasses, and thornywoodlands (Chundawat et al, 1997). Thecharacteristic floral species of this area includestree species such as Tectona grandis, Diospyrosmelanoxylon, Madhuca indica, Buchnania latifolia,Anogeissus latifolia, A. pendula, Lanneacoromandelica, Boswelia serrata, etc. Majorshrub species include Lantana camera, Grewiasp., Nyctanthus arbortristis, Ixora sp., Zyziphusmauritiana, Z. oenoplea, etc. The dominant grassspecies are Apluda mutica, Themeda quadrivalvis,

Heteropogon contortus, Aristida sp., Eragrostissp., etc.

The park supports a diverse fauna.Among the large predators found in the area aretiger (Panthera tigris tigris), leopard (Pantherapardus), sloth bear (Melursus ursinus), wild dogs(Cuon alpinus) and wolf (Canis lupus laniger). Theherbivores include, sambar (Cervus unicolor) ,chital (Axis axis), nilgai (Boselaphustragocamelus), chinkara (Gazella gazella) andchausinga (Tetraceros quadricornis). Othermammalian species commonly found are hyaena(Hyaena hyaena), jungle cat (Felis chuas), wildpig (Sus scrofa), Hanuman langur(Semnopithecus entellus), jackal (Canis aureius),Indian fox (Vulpes bengalensis) and Indian treeshrew (Anathana ellioti). The avifauna of the parkis rich and is represented by over 200 speciesof birds.

There are 13 villages inside the NationalPark with a total human and cattle populationof approximately 2,500 and 9,500 respectively.The main occupation of people residing inside thepark is agriculture and cattle rearing.

8

Panna Tiger Reserve (543 kmPanna Tiger Reserve (543 km22))Provides a representative site in a Dry Tropical Forest.Provides a representative site in a Dry Tropical Forest.Study attempts to find out the threat responsible for the declineStudy attempts to find out the threat responsible for the declineof tiger population in Dry Forest habitats where it is most vulnerable.of tiger population in Dry Forest habitats where it is most vulnerable.

Panna Ti

ger Rese

rve (543

km

Panna Ti

ger Rese

rve (543

km22))

Figure 1.. Map of Panna NP and its adjoining area.

General Methods

A preliminary survey was carried outtowards the end of 1995 in Panna National Parkto determine the suitability of the area forconducting a radio-telemetry project. Fieldworkbegan in October 1996, during which researchactivities such as the recording of tigermovement with the help of radio telemetry,estimation of tiger prey abundance, habitatevaluation, collection of scats to determine thefood habits of tiger and determination of bioticpressures on the tiger and its prey wereundertaken. The study’s research workedcontinued until December 2004.

Estimation of Prey Population Prey abundance was estimated by distancesampling (Burnham et al. 1980; Buckland et al.1993). Distance sampling through line transectsurveys has been used very widely andsuccessfully in tropical forests of the sub-continent (Karanth and Sunquist, 1992;Khan etal., 1996; Biswas and Sanker, 2002; Jathannaet. al. 2002; Karanth et. al., 2002; and Bagchiet al., 2003). For this purpose, eleven straighttransect lines were established in different

strata identified within the intensive study areaand marked at regular intervals(Figure 2). Thesetransect lines were maintained every year andwalked regularly to gather information on groupsize and population structure and to estimatethe abundance of tiger prey species. A similar exercise was also conducted usingvehicles and the roads were used as lines; thesewere monitored regularly, twice or thrice everymonth. Information on abundance, distribution,herd size and population structure of ungulatesover a larger area of the Park was collected fromthis field exercise. For this method, the existingroad network of the Park was used. ‘Distance’(Laake et al. 1994), a computer softwareprogramme, was used for the analysis of theabove data.

Home Range and Movement PatternRadio-collaring and monitoring of animals

In order to collect information on homerange the movement, habitat use and activityof tigers and their major prey, several animalswere radio-collared. Permission for chemicalcapture and radio-collaring of tigers and their

9

prey ? sambar, chital and nilgai ? was obtainedfrom the Government of Madhya Pradesh andthe Ministry of Environment and Forests,Government of India. In all, seven tigers (severalof the tigers were collared more than once overthe years), five sambar and two chital werechemically immobilized and radio-collared. All thechemical restraint operations were conductedby professional veterinary experts who came fromthe Wildlife Institute of India, from the Universityof Hissar (anaesthesia specialist) and from theVeterinary college, Jabalpur.

Chemical Restraints of TigersFor radio-collaring purposes it is necessary

to chemically restrain the tigers. First thetargeted tiger was located by the research team;then a team of veterinary professionals, whichincluded a wildlife veterinary expert, anaesthesiaexpert and another veterinary doctor, cautiouslyapproached the animal on elephant back. In thecase of collared animals (for replacing the oldfunctional collar) only one elephant was used.But in the case of a new individual or whenreplacing a non-functional collar, two elephantswere used and a few observers were place on atree at two to three vantage points to keep aneye on the tiger’s movements after the drug wasinjected. After locating the tiger, the elephantwith the veterinary doctor approached the

animal within 25-30 metres and waited until theanimal provided a good view for darting. The research team used Teleinject equipmentfor darting the animal. A new but very effectiveand safe drug called Meditomedine (trade name:Zalopine) in combination with Ketamin was usedto capture the animal and “Antisedan” was usedto reverse the effect of the tranquilization. Onone occasion the research team also used thecombination Rompun and Ketamin for chemicalcapture. It was ensured that pressure on theprojectile was kept at the minimum required inorder to minimize the impact of the dart as ithit the animal. This is important to reducetrauma otherwise caused by the impact of thedart on the animal. We were able to reduce theimpact of the dart to its bare minimum so thaton most occasions the tigers did not move morethen thirty to forty metres after contact. Insome cases the tiger moved even less then tenmetres after it had been darted. This allowed the research team and theveterinary doctors to monitor the animalthroughout the induction time of the drug. Afterten to twelve minutes, when the animal’s headwas rested on the ground, it was approachedfrom behind. The animal’s reflexes were firsttested with the help of a long stick and laterthe tail was touched to assess the effect of the

10

drug on the animal. Once the tiger’s status wasproperly assessed, two or three members of theresearch team quickly moved in to fix the radio-collar on the animal and take the morphometricmeasurements. On average, this took less thentwenty minutes. As soon as the collar was placedand measurements recorded, some photographswere taken for identification purposes. After theresearch team moved away from the animal, adose of antidote was given intra-muscularly attwo to three places. The recovery of thetranquilized animal was monitored from elephantback. Recovery of tiger usually took ten to fifteenminutes when using Meditomedine and longerwhen the combination of Xylezine and Ketaminewas used. Recovery in the case of Meditomedinewas smooth and quick and the animal walkedaway with coordinated steps within twentyminutes after the antidote was injected. Theresearch team continued to monitor the animalintermittently for the next eight to ten hours.

Chemical Restraint of Prey SpeciesTiger’s major prey, sambar and chital, were

captured using the same drugs and equipment,that were used for tiger. Animals were

approached from an open vehicle and darted froma distance of 25-30 metres and also from hides.After the darting, the animal was observed froma distance and its activities were monitored withthe help of binoculars. On average the dartedanimal moved 50-60 metres from the site it wasdarted and it took 6-8 minutes for it to go down.Once the animal went down, the research teamwaited for twelve to fifteen minutes beforecautiously approaching it. From ten metres, theanimal’s status was again assessed before itwas approached carefully and touched.Immediately a dark cloth was used to cover theeyes and a radio-collar was attached. Afterrecording the measurements, the antidote wasgiven and recovery of the animal was observedfrom 20-30-metres distance. The animals tookfifteen to twenty minutes to be able to walkproperly. These animals were monitored closelyby the research team for the next 24 hours tominimize the possibility of the predation.

Monitoring of Radio-collared AnimalsRadio-collared animals were tracked on a

systematic schedule to obtain the locations ofthe animal. The locations are determined by

triangulation method (Tester and Sniff1957, Heezen and Tester 1967 & Springer1979) and wherever possible by homingtechnique (Mech 1983). For this all roadswere marked at 500m intervals and in someplaces every 250m. UTM coordinates foreach of these points were recorded with thehelp of a GPS. In addition to these,coordinates of several vantage locationswere noted down. Once the radio-collaredanimal is contacted, compass bearings aretaken and recorded from these knownpoints with the help of directional antenna.This exercise was repeated from three tofour known points. Later with the help ofthe computer software “LOAS” the animal’slocation is determined using the UTMcoordinates and compass bearings. Theselocations of the animal are used fordetermining home ranges, movementpattern and habitat use, by the radio-collared animals.Locations for every animal were obtained

twice or thrice a week to determine the

11

Tr-9Tr -4

Tr-3

Tr-2

Tr -6Tr-7

Tr-5

Tr-8

Tr-11

Tr -1

Tr-10

Panna National ParkDistribution of Transect lines in the Study Area

Figure 2.. Park of the Panna NP map showing transect lines

home ranges and habitat use pattern over theseasons. These animals were also monitoredcontinuously for a twenty-four hour period oncea month, to gather information on the dailymovement and habitat use on a daily basis. Forthe twenty-four hours monitoring, locations areobtained every hour for tiger and every two hoursfor prey species and activity is recorded everyfifteen minutes.

Habitat Use by Tiger A detailed habitat map was prepared usingsatellite LISS-III data. Seven major habitatclasses were identified. This habitatclassification was verified andmodified after ground truthing.Based on this information, a detailedhabitat map was prepared for thePanna National Park and its adjoiningtiger habitats (Figure 1). The final mapwas used to determine habitat use bythe tigers and their prey species byplotting their locations on the mapand calculating the frequency ofoccurrence in different habitat types.This exercise was repeated fordifferent seasons to determineseasonal habitat use. Using this habitat map of Panna

Tiger Reserve and other layers such as grazingpressures, village locations, water distributionand other anthropogenic pressures and densityestimation of major prey species in differenthabitats, predictive models for the distributionof major prey species of tiger, sambar, chital andnilgai were developed using GIS. These maps wereused to determine the distribution of tiger inrelation to the availability of its major preyspecies. In addition, to supplement this information,throughout the intensive study areas, ‘controlplots’ were established on a grid basis every one-minute. These plots were later monitored at the

Figure 3. Research team radio-collaring a male sambar

12

end of every season for tree and shrub density,canopy and grass cover, stalking and hiding cover,grass height, disturbance, abundance of tigerprey by indirect evidences (Muller-Dombois &Ellenberg 1974) and a temperature data loggerwas placed at selected plots to record thetemperature every hour. This providedinformation on the availability of resources forthe animals. Use of these resources by theseanimals was determined by visiting the locationsobtained through radio-tracking of animals andconducting similar habitat evaluations for thelocations of each radio-collared animal. Tominimize the workload, instead of visiting all thelocations, 40% of the locations were selected,randomly, to determine the habitat use by theseanimals.

Food Habits and Predation by TigersFood habits of tiger are determined by

identifying the prey remains in scats andinformation collected from kill sites (Schaller1967, Johnsingh 1983, Karanth 1993, Chellam1993). In the field it is not always possible tofind each and every kill made by the tiger.Therefore, using only found kills leads tounavoidably biased information and under-represents the utilization of smaller prey species,which are consumed relatively fast (Sunquist1981, Johnsingh 1983, Karanth & Sunquist1995). Therefore, to have a reliable picture of foodhabits of tigers, scats were collected from thestudy area and analyzed. Prey remains in thescats of the carnivores are mostly the hairs andpieces of bones: these remain undigested whilepassing through the gut (Sunquist 1981;Karanth & Sunquist 1995; Mukherjee et al.1994). Through microhistological study of boneand hair, prey species were identified (Koppikarand Sabnis ???).

TIGER PREY-I: Group dynamics, population structure andestimation of animal and biomass density. Tiger has a very wide distribution in Asia andoccupies variety of habitats throughout itsrange (Sunquist et al., 1999). Here it predatesmainly on large ungulates of tropical andtemperate forests (Schaller, 1967; Sunquist,1981; Johnsingh, 1983; Seidensticker andMcDougal, 1993; Karanth and Sunquist, 1995;Biswas and Sanker, 2002, Bagchi et. al., 2003).Like other large carnivores, its abundance isgoverned by the availability of its prey (Sunquistand Sunquist, 1989). Tiger has evolved as aspecialist predator of large deer, wild cattle andpigs and its present distribution is limited tohabitats where populations of its major prey stillsurvive in abundance (Sunquist et al., 1999).Recent studies indicate that loss of prey is oneof the major factors affecting the viability of thetiger populations (Karanth and Stith, 1999) andindicate that a demographically viable tigerpopulation can be protected provided that theirprey base is managed intensively. Therefore, inorder to save wild tigers, effective managementof large herbivore populations is an importantconservation issue in tiger habitats. Despite theconservation significance of Tropical Dry Forests

for tiger and other co-predators, very littlequantitative information is available from thisecosystem. This largest tiger habitat is highlyfragmented (Wikramanayake, et al., 1998) andsuffers from forest fires, poaching, competitionwith livestock and loss of habitat and habitatquality (Panwar, 1987; Karanth 1991; Deb Roy,1996; Seidensticker, 1997, Chundawat andGogate, 2001). For effective conservationmeasures to save tigers in Dry Forests, intensivemanagement of prey populations requirestrategies to manage prey in densities that cansupport a demographically viable tigerpopulation. Basic quantitative information onprey availability for tigers is required for this.

MethodsPopulation structure and Group size The prey population of Panna Tiger Reservewas monitored regularly on a very systematicschedule by counting animals from the vehicle.The intensive study area included most of theforests of the three ranges of Panna, Hinautaand Madla. The forest roads of the Reserve inthese ranges were used as transect lines and

13

monitored twice or thrice every monththroughout the year. This was not possible duringthe monsoon months, due to heavy rain and badroad conditions, but even then the roadtransects were conducted at least once amonth. Whenever an animal was seen,information on the species, number, group size,age and sex and the perpendicular distance fromthe road was recorded, in addition to the dateand time, activity and transect number. Fromthis data set, information on demographicparameters of the prey population is generated.

Estimation prey population In this study we used distance sampling byline transect to estimate abundance of tiger preyin the Tropical Dry Forest of Panna Tiger Reserve.Line transect surveys were conducted in years:2000, 2001 and 2003. For these surveys, nineline transects of variable length, ranging from1.93-2.85 km, were marked. They were alsoprepared for better view on the line and moresilent walk. In 2000, these nine lines were walked in themorning twice every month from October to June.Later this approach was modified and in 2001,all the lines were walked simultaneously everymorning and evening for fifteen days in March.Further modification was made in the 2003survey: only three lines were walkedsimultaneously morning and evening and thesewere repeated every fourth day in February andMarch. Thus in total 24 (12 days x 2 times aday) temporal replicate surveys for each line wereconducted. Data from these three surveys areused here to discuss the best design approachesand appropriate season for surveys in tropicalforests where visibility changes considerably withthe season and can thereby affect densityestimates. These data sets are also analyzedto determine the efficiency of the survey withoutcompromising on most of the assumptions. Weseek to use the results to help in designing futuresurveys that will achieve the desired objectivesmost effectively. Densities and biomass densities of tiger preyspecies were calculated from the data obtainedfrom the most recent, 2003, survey; this is usedfor comparisons with estimates obtained fromother habitats of the Indian sub-continent. For

every group sighting, information on species, thenumber of individuals, radial distance (with helpof laser range finder) and radial angle wererecorded. For analysis of data, the computerprogram DISTANCE 4.0 (Thomas et al., 2002-b) was used.

Analysis For density estimates and related analysesconcerning variance, the nine transect lines wereconsidered as spatial replicates and all thetemporal replications on particular transectlines were pooled. We followed the standardprocedures for analysis detailed in Buckland etal. (2001) and Thomas et al. (2002b). Clustersizes were estimated using size bias regressionmethod by regressing the natural log of clustersize against estimated probability of detectionat distance x, g(x). Whenever the regression wasnot found significant at alpha level of 0.15, meanof cluster sizes was used. Both manual as wellas automatic pooling functions were used inDistance 4.0 to fit the curve at the lowestpossible value of ?? (Chi-square) To compare seasonal and morning-eveningestimates obtained by distance software, onefactor ANOVA was used with available estimatesand respective standard deviations (Zar, 1984).Changes in other parameters like seasonal groupsize and radial distance were tested using onefactor ANOVA. Trends were tested for accuracywith the help of maximum R2 value and withstudent’s t-test for testing the significance ofa regression.

ResultsPopulation structure and Group size In this report information on populationstructure and group size of tiger prey is reportedfrom the most recent 2003 monthly surveys.Prey species include sambar, chital, nilgai, four-horned antelope, chinkara and wild pig. A totalof 4,434 animals were sighted of which 3,332animals were aged and sexed. These included 252chinkara, 1,278 chital, 125 four-horned antelope,1,409 nilgai, 794 sambar and 134 wild pig. Theaverage group size observed during the study is:sambar 3 (SE ± 0.14), chital 5 (SE ± 0.41), nilgai2.8 (SE ± 0.11), four-horned antelope 1.3 (SE ±0.06), chinkara 1.68 (SE ± 0.07) and wild pig4.2 (SE ± 0.4). Except for chital and wild pig, no

14

significant difference in group size is observedfor the different seasons in most prey species(Figure 4). Both chital and wild pigs are seen inlarger groups during the monsoon months thanin summer and winter. Group size range observedis 1-20 for sambar, 1-64 for chital, 1-19 for nilgai,1-5 for four-horned antelope, 1-6 for chinkara and1-14 for wild pig. Details of population structure for preyspecies is given in figure 6. No change in thepopulation structure (age and sex ratio) isobserved from the previously reported data(Chundawat, 2001). Chital to fawn ratio remainslow at 24 fawns for every 100 females. In thecase of wild pig, high female to young ratio wasseen and it was 100:100, but low female toyearling ratio 100:22.9 indicates high mortalityat an early age. Male to female ratio remainedbiased towards female in all the prey speciesobserved (Table 1).

Line transect samplingIn this report, data from 2000, 2001 and 2003surveys is analyzed and reported. In the monthlysurvey of 2000, the total length walked was233.6 km and 339 groups of animal weresighted. In the 2001 and 2003 surveys, the totallength walked was 520.7 km and 466.06 kmrespectively. The total number of animal groupsseen was 1,014 during the 2001 survey and 1,023groups in the 2003 survey. We pooled the 2000distance sampling data bi-monthly to analyzeand detect changes in encounter rates, groupsizes and sighting distances and their likelyeffect on density estimates. Encounter rates ofall the prey species varied significantly betweenthe two months groups (Chital2000: F=3.538,p<0.05; Nilgai2000: F=7.494, p<0.001;Sambar2000: F=5.049, p<0.005). Thecumulative encounter rates of the prey speciesof tigers found in Panna Tiger Reserve reachhighest during the months of February andMarch (Figure 5). Ungulate density variedsignificantly (Chital2000: F=3.066, p<0.05;Nilgai2000: F=2.87; p<0.05; Sambar2000:F=4.031, p<0.01).

The subsequent two surveys wereconducted during February and March. Duringthe 2001 and 2003 surveys, a larger distancewas covered in a relatively short duration by

walking all the transect lines, simultaneously,every day in the morning and evening. The surveysin February and March resulted in a largerdetection of animal groups for all the speciesthan observed in the previous monthly surveys. Mean group size for most of the species didnot vary significantly during the 2001 (fifteendays) and 2003 (forty five days) surveys. Radialdistances of chital changed significantly in 2001(F=6.68, p=0.005) as well as in 2003(F=5.284, p=0.000). This was primarily due tofire on the transect lines towards the end of boththe surveys. These fires opened the habitat andthis increase in the visibility may be the majoraffecting factor that brought the significantchanges in radial distances encountered. But,during these surveys no significant change in theencounter rate was observed for most of the preyspecies except for nilgai, where it showed agradual increase from 0.3 to 0.9 per km in the2003 survey. Encounter rates for most species in themorning and evening surveys of 2001 and 2003differed significantly (Sambar2003: F=5.59p=0.05; Nilgai2003: F=11.35, p=0.005;Sambar2001: F=9.24, p=0.01) and these rateswere consistently lower in the evening for all thespecies. Thus there is a significant effect on thedensity estimate for a few of the species whenthe morning and evening data is pooled. (onetailed tests Nilgai2003: F = 16.63, p=0.001;Sambar2001: F=11.03, p=0.005). In the 2003survey, we walked these lines in the morning andevening every fourth day to examine the effectof disturbance from repeated walking but theresults were similar with a lower encounter ratein the evening walks. It was also noticed that generally for all thespecies except sambar the encounter rate, inmorning and evening, gradually declined with thelength of time spent in walking the transectlength. In the case of chital and nilgai it declinedsignificantly. There was a significant negativetrend in case of chital in both years where theencounter rate dropped on the morning(R22001=0.73; t2001=3.28, p=0.05,R22003=0.76; t2003=4.33, p=0.005) as wellas evening (R22001=0.88; t2001=5.43, p=0.01;R22003=0.88; t2003=6.53, p=0.001)transects. Similarly for nilgai it showed a

15

significant declining trend in year 2003(R22001=0.48; t2001=1.91, p=0.2;R22003=0.58; t2003=2.87, p=0.05) in themorning and in the evening in 2001(R22001=0.93; t=7.18, p=0.002;R22003=0.26; t=1.47, p=0.2). However,although the sambar encounter rate declinedsimilarly in the morning, on the evening transectthe encounter rates marginally increased closeto dusk (Figure 7). This can be explained by theactivity pattern determined from the radio-collared animals (Figure 8).

Density and Biomass estimation Density of tiger prey is estimated from themost recent count conducted in 2003. Thepooled prey abundance is estimated to be 46.32prey/km2, (excluding langur). Details of theestimated prey densities and other relevantparameters are described in Table 3. As in mostof the tiger habitats of the subcontinent, chital,nilgai and sambar dominate the wild tiger preypopulation in Panna Tiger Reserve. In terms ofnumber of animals per square kilometre, chitalis the most abundant animal. In the Reservethe contribution of smaller prey (<25 kg andincluding langur) is 30%, medium sized prey (25- <55kg), which includes chital and wild pig is38% and large prey (>55 kg), which includessambar and nilgai, is 31%. Most frequentlyencountered prey in the Reserve is nilgai followedby sambar and chital. The ungulate prey biomass densityestimated for the Tropical Dry Forest of PannaTiger Reserve (4,057 kg/km2) is on the high sidefor the subcontinent. But, unusually for the sub-continent, most of the biomass (over 70%) iscontributed by two prey species, nilgai andsambar. The contribution of chital, the tiger’smost common and major prey throughout thesubcontinent, is only 19.69%. Contribution fromother smaller prey species is minimal (< 7%).

DiscussionGroup size and Population structure It is observed in Panna Tiger Reserve thatgroup size of most of the prey species is relativelysmall (Karanth and Sunquist, 1992; Khan et al.,1996; Biswas and Sanker, 2002; Jathanna et.al. 2002; Karanth et. al., 2002; and Bagchi etal., 2003). It is more pronounced in chital

population, though a comparison of group sizebetween 2001 and 2003 counts shows amarginal increase in group sizes of chital to amean of 5 (± 0.41) from a mean group size of 3.Group size could be an important indicator ofpoor resource availability and quality and habitatsuitability limiting the reproductive success ofthe population (Jarman, 1974). Smaller groupsize in tiger prey population is an importantfinding for Tropical Forest, which is the largesttiger habitat in the sub-continent and needsfurther investigation to ascertain the factorresponsible for affecting the group size in thistype of habitat. Most of the prey species, other than chitaland wild pig, did not formed larger groups indifferent seasons. In the chital and wild pigpopulations, group size was larger during themonsoon months, when forage is abundant.Smaller groups were seen in winter, amongst thesambar, chital and wild pig. We observed nochange across the seasons in the sex and ageratio of all the prey populations. It is also observed that the female to fawnratio in chital is very low in comparison to otherchital populations from different but moreproductive tiger habitats (Mishra, 1982;Karanth, 1993; Khan et al., 1989, Kumar, 2000).It is also noticeable that the survival of thispopulation of fawns into the next age class ofyearling is also very low. This indicates poorreproduction in the chital population and highmortality in the yearling age class. Panna, whichis characterised as Dry Tropical Forest, hasextensive open miscellaneous forest with densegrass cover. This does provide suitable habitatsfor chital with high abundance of graze. Grassgrowth stops soon after it flowers and seeds,as a result of the long dry spell soon after themonsoon. After December, the dead, above-ground biomass of grass provides very poorquality forage in terms of nutrition. This is at atime when the chital population requires qualitynutrition prior to their readiness for rut in Marchand April. This could be one of the reasons forlow fawn ratio in Panna. A similar high mortality is also noticed inthe sambar population from a healthy >40fawns/100 females to <20. This, as mentionedin the earlier reports, is an important aspect

16

10022.956.25100Wildpig

9.42.872100Chinkara

302.845.71004horn

35.9820.7951100Nilgai

41.416.9433.89100Sambar

24.2710.7541100Chital

FawnYearlingMaleFemaleSpecies

10022.956.25100Wildpig

9.42.872100Chinkara

302.845.71004horn

35.9820.7951100Nilgai

41.416.9433.89100Sambar

24.2710.7541100Chital

FawnYearlingMaleFemaleSpecies

for the management to look into for the bettermanagement of prey population especiallysambar which is a preferred prey species of tigerin the subcontinent (Biswas and Sanker, 2002;Karanth et. al., 2002; and Bagchi et al., 2003).A high female to young ratio was observed in thewild pig population but its female to yearling ratiois low, suggesting high mortality at this agecategory. This could be due to high predation onpiglets by predators such tiger, leopard andjackal. Sambar, four horned antelope, and chinkaraare mostly seen singly or in small family groups,which are comprised of mother, fawn and yearling.This information is further supplemented by theobservations made on the radio-collared femalesof sambar. Males are usually solitary exceptduring the rut, when they associate with femalegroups or join other males temporarily after therut is over. Female family groups occasionallyassociate with other such groups for a shorttime at their favourite feeding places. In contrastto these observations made on the three preyspecies above, chital and nilgai are morefrequently seen in larger groups or associations,indicating their social nature.

Estimation prey population The density estimate of 46.32 preys/km2 inPanna Tiger Reserve is fairly high for a TropicalDry Forest of the subcontinent (Karanth andSunquist, 1992; Khan et al., 1996; Biswas andSanker, 2002; Bagchi et al., 2003). Because

mega-herbivores suchas wild buffaloes(Bubalus bubalis),rhinos (Rhinocerosunicornis) andelephants (Elaphusmaximus), whichcontribute very littleto the diet of tiger, areabsent from a largepart of the TropicalDry Forest, they areexcluded forcomparison of theprey populations fromimportant tigerhabitats. Biomass

densities from these tiger habitats indicatethat, on average, medium sized prey (46.64 %)and large prey (52.27 %) contribute almostequally to the prey availability. But interestingly,in terms of number of animals, medium sized prey- such as chital, wild pig and hog deer –contribute, on average, over 70 % of preyavailability (Table 4). In contrast to theseprotected habitats, in the Panna Tiger Reserve,the contribution by chital and other medium preyto the prey population is small, both in term ofbiomass density (21.26 %) and the number ofanimals (39.24%). The low abundance of chital, one of the tiger’smajor prey species, could be an important factorto the tiger ecology in Panna. Being the onlygrazing wild herbivore (Eisenberg andSeidensticker, 1976; Putman, 1988, Khan et al.,1996), chital is particularly and highlysusceptible to grazing pressure of domesticlivestock populations (Khan, 1996, Jathana etal., 2003). A large population of over 9000domestic cattle within Panna Tiger Reserve andextensive accidental fires in the past could bethe reasons that limit the chital population here.Despite their low nutritive value, chital prefergrasses even during the dry summer months(Khan et al., 1996). Dry season fires aredevastating in Panna, completely eradicatinggrasses from the burnt areas. In the recentpast, rehabilitation of three villages and intensivefire protection measures have created extensivegrasslands and restored them as suitable

Table 1. Sex ratios of tiger prey described animals per hundred females in PannaTiger Reserve.

17

C h e e t a l

0

2

4

6

8

10

m o n s n summr w i n t r Annua l

Me

an

gro

up

siz

e

C h i n k a r a

0

0.5

1

1.5

2

monsn summr win t r A n n u a l

Me

an

gro

up

siz

e

F o u r H o r n A n t e l o p e

0

0 . 5

1

1 . 5

2

m o n s n s u m m r wint r a n n u a l

Me

an

gro

up

siz

e

N i l g a i

0

0.5

1

1.5

2

2.5

3

m o n s n s u m m r wintr a n n u a l

Me

an

gro

up

siz

e

S a m b a r

0

1

2

3

4


Me

an

gro

up

siz

e

W i l d b o a r

012

34567

m o n s n s u m m r w i n t r a n n u a l

Me

an

gro

up

siz

e

C h e e t a l

0

2

4

6

8

10

m o n s n summr w i n t r Annua l

Me

an

gro

up

siz

e

C h i n k a r a

0

0.5

1

1.5

2

monsn summr win t r A n n u a l

Me

an

gro

up

siz

e

F o u r H o r n A n t e l o p e

0

0 . 5

1

1 . 5

2


Me

an

gro

up

siz

e

N i l g a i

0

0.5

1

1.5

2

2.5

3

m o n s n s u m m r wintr a n n u a l

Me

an

gro

up

siz

e

S a m b a r

0

1

2

3

4


Me

an

gro

up

siz

e

W i l d b o a r

012

34567

m o n s n s u m m r w i n t r a n n u a l

Me

an

gro

up

siz

e

30.859.709.41-508.01197Langur

0038.161.81-41.688Chinkara

002.460.91-123.4143Wild pig

0017.989.01-41.2144Fourhorn

2.421.940.435.11-172.8909Nilgai

6.927.935.329.61-443.96963Chital

017.743.039.11-102.2577Sambar

>104-102-31

Percent occurrence in size class sizeRangeMeanGroup size

AnimalSighted

Species

30.859.709.41-508.01197Langur

0038.161.81-41.688Chinkara

002.460.91-123.4143Wild pig

0017.989.01-41.2144Fourhorn

2.421.940.435.11-172.8909Nilgai

6.927.935.329.61-443.96963Chital

017.743.039.11-102.2577Sambar

>104-102-31

Percent occurrence in size class sizeRangeMeanGroup size

AnimalSighted

Species

habitats for chital. As a result the chitalpopulation is recovering, mainly in areas recentlymade free from human pressure, as has beendocumented in other protected forests of thesubcontinent (Karanth and Sunquist, 1992;Khan et al., 1996). Density estimates of tiger prey from PenchNational, Ranthambhore National Park, the Girforest of Gujarat and Panna Tiger Reserve

indicate that Tropical Dry Forests - the largesttiger habitat in India (Wikramanayake, et al.,1999) - can support fairly high prey biomass,provided the habitat is adequately protected(Eisenberg and Seidensticker, 1976; Karanth andSunquist, 1992; Khan et al., 1996; Karanth andNicholas, 1998; Biswas and Sanker, 2002). TheDry forests of Panna support ungulatepopulations which have evolved in different

Figure 4. Mean group size of prey species in Panna Tiger Reserve

Table 2. Mean group sizes, range and frequency of group size of tiger prey in Panna Tiger Reserve.

18

0

0.2

0.4

0.6

0.8

1

1.2

1.4

1.6

1.8

2

Oct_Nov Dec_Jan Feb_Mar Apr_May Jun

Enc

ount

er r

ate

Chital Nilgai Sambar All Ungl.

environments – one in open habitats and theother in forest mosaic. Nilgai and chinkararepresent a prey population partial to openhabitats, whereas chital and sambar are forestand forest-edge dwellers ((Schaller, 1967; Prater,1988; Eisenberg and Seidensticker, 1977).Because tiger has evolved as a specializedforest-edge predator following the cervidradiation in Asia (Sunquist, et. al., 1999), itssurvival and hunting strategies are more cuedto cervids than other prey species occupying theopen habitats such as nilgai. Therefore in DryForests, despite high abundance of prey such asnilgai in open habitats, this does not necessarilytranslate entirely into tiger prey availability. In tropical forests of the subcontinent,ecological conditions change drastically overtime and space and this can affect animaldistribution and detection (Thomas et al.,2002). Soon after the monsoon, visibility inforest habitats gradually increases with loss offoliage as the long dry season progresses. Lossof foliage not only makes it easier to detectanimals but they can now be sighted from longerdistances, significantly increasing the radialdistances. In addition to this, the significantchange in group size over the seasons, not onlyaffects the number of animals encountered butcan affect the encounter rate of the groups.Moreover, during peak summer months whenwater is available only at a few sites, most ofthe ungulates congregate near these habitats.In these changing ecological circumstances,monthly surveys do not provide a reliable way toestimate ungulate densities in a Tropical Dry

Forest such as the Panna Tiger Reserve.Pooling data from the surveys conducted overan extended period time of several months aretherefore likely to be prone to limitations suchas those encountered during the monthlysurveys of 2000 in Panna TR. It is clear from the monthly surveys in2000 that the highest encounter rates ofdetection for most ungulate prey are obtainedduring February and March. Moreover, groupsizes during these months are also closer tothe mean estimated for the year. Therefore,for estimation of wild herbivore populations ina Tropical Dry Forest, February/March appearsto be the most suitable time. Walking transect

lines simultaneously and repeatedly everymorning and evening maximized the efficiency ofthe surveys (Karanth and Nicholas, 1998). Byconducting these surveys within a short timespan (ranging from 15 to 40 days), the errorassociated with varying encounter rates, changein group sizes and effect of visibility on radialdistance and ESW, is minimized effectively in thisstudy. But the problem of low encounter ratesduring the evening count on the same line affectsthe density estimation. One of the likely reasonsfor low density estimates in the evening couldbe the fact that during these intensive surveys,all the lines were walked repeatedly in the morningand evening every day. It is speculated that thismay have caused enough disturbance for theanimals to move away from the area and affectthe low evening encounter rates. Another factor could be the activity patternof prey species. It is generally assumed thatmost of the animals are active during the earlymorning and evening (Karanth and Sunquist,1992). The radio telemetry study of sambar andchital in Panna indicates that the activitypattern of herbivores is a likely factor resultingin the low detection and density estimate in theevening. The first half of the morning surveyscoincide with the peak activity period, whichdeclines gradually. Similarly, increasing encounterrates in the case of sambar towards the end ofthe evening coincide with its increase in activity.But decreasing encounter rates towards theevening for chital and nilgai is difficult to explainwith the existing data set. The effect of repeated

Figure 5. Bi-monthly (pooled) encounter rates of majorprey species of tiger during the 2000 survey in PNP

19

Fm-A

D

Fm-SA

Male-1

Male-2

Yrlng

Fawn

0

10

20

30

40

50Sambar monsn

summr

wintr

Fm-AD

Fm-SA

Male-1

Male-2

Yrlng

Fawn

0

10

20

30

40

50

Nilgai monsn

summr

wintr

Fm-A

D

Fm-S

A

Male-1

Male-2

Yrlng

Fawn

0

10

20

30

40

50Wild boar monsn

summr

wintr

Fm-A

D

Fm-SA

Male-1

Male-2

Yrlng

Fawn

0

10

20

30

40

50

Cheetal wintr

summr

monsn

Fm-AD

Fm-S

A

Male-1

Male-2

YrlngFa

wn

0

1020

30

40

50

60

Chinkara monsn

summr

wintr

Fm-AD

Fm-SA

Male-1

Male-2

YrlngFa

wn

010

20

30

40

50

60

Four horn monsn

summr

wintr

Fm-A

D

Fm-SA

Male-1

Male-2

Yrlng

Fawn

0

10

20

30

40

50Sambar monsn

summr

wintr

Fm-A

D

Fm-SA

Male-1

Male-2

Yrlng

Fawn

0

10

20

30

40

50Sambar monsn

summr

wintr

Fm-AD

Fm-SA

Male-1

Male-2

Yrlng

Fawn

0

10

20

30

40

50

Nilgai monsn

summr

wintr

Fm-AD

Fm-SA

Male-1

Male-2

Yrlng

Fawn

0

10

20

30

40

50

Nilgai monsn

summr

wintr

Fm-A

D

Fm-S

A

Male-1

Male-2

Yrlng

Fawn

0

10

20

30

40

50Wild boar monsn

summr

wintr

Fm-A

D

Fm-S

A

Male-1

Male-2

Yrlng

Fawn

0

10

20

30

40

50Wild boar monsn

summr

wintr

Fm-A

D

Fm-SA

Male-1

Male-2

Yrlng

Fawn

0

10

20

30

40

50

Cheetal wintr

summr

monsn

Fm-A

D

Fm-SA

Male-1

Male-2

Yrlng

Fawn

0

10

20

30

40

50

Cheetal wintr

summr

monsn

Fm-AD

Fm-S

A

Male-1

Male-2

YrlngFa

wn

0

1020

30

40

50

60

Chinkara monsn

summr

wintr

Fm-AD

Fm-S

A

Male-1

Male-2

YrlngFa

wn

0

1020

30

40

50

60

Chinkara monsn

summr

wintr

Fm-AD

Fm-SA

Male-1

Male-2

YrlngFa

wn

010

20

30

40

50

60

Four horn monsn

summr

wintr

Fm-AD

Fm-SA

Male-1

Male-2

YrlngFa

wn

010

20

30

40

50

60

Four horn monsn

summr

wintr

walks on the encounter rate can be tested byconducting a similar experimental exercise wherelines are walked once only in a day to see theeffect of disturbance caused by the previouswalk. In the present data set this variation inthe evening and morning count has possibly

Figure 6. Population structure of prey species (in percentage) found in PNP.

caused under estimation of prey population. These trends were consistent during all thesurveys and highlight an additional point thatcounting animals for more then two hours canhave a significant effect on the density estimatebecause of the shift in activity pattern. This

20

125.1

816.6

1.2

011.5

50.1

320.4

19.4

70.1

813.9

065

2003

186.2

025.6

2.5

18.2

20.2

836.4

111

.726.7

20.6

8130

2001

227.8

9

36.4

2.2

511.2

30.1

758.5

710.9

37.8

25.3

12000

Lan

gur

63.3

245.1

1.0

91.8

10.1

75.3

20.

7345.5

1.9

785

2003

29.2

141.6

0.7

21.6

20.1

2.1

80.3

842.1

0.9

144

2001

73.6

3

32

49.4

1.2

21.8

80.1

6.9

40.

7650.3

2.3

025

2000

Ch

inkar

a

30.2

252.1

0.3

74.0

50.0

44.6

20.

4957.8

1.5

120

2003

27.3

442.6

0.5

83.2

80.0

83.4

40.

5446.5

1.3

629

2001

NA

20

Da

tase

t to

o sm

all

to e

stim

ate

den

sity

62000

Wild

pig

56.7

42

20.2

2.2

11.2

20.1

74.1

91.

7320.6

2.7

082

2003

82.9

919

3.3

21.2

30.2

15.9

52.6

219.2

3.9

596

2001

62.4

75

21

29.5

1.8

41.6

10.1

35.7

11.

531.9

2.9

731

2000

4H

An

telo

pe

1639

24.5

4.4

02.9

80.6

823.0

97.

4426

13.1

1333

2003

787.6

237

2.6

72.7

20.6

13.9

92.8

337.2

6.3

0126

2001

1421.7

5

125

26

3.6

23.1

40.4

421.0

66.

1429.1

11.3

7107

2000

Nilg

ai

1387.5

733.7

3.9

52.6

20.4

522.0

04.

8734.1

10.3

5223

2003

1175.5

836.9

3.9

72.

30.4

719.4

13.9

637.2

8.7

7234

2001

2510.8

9

134

34.2

7.1

82.6

10.6

41.5

28.

4534.9

18.7

3147

2000

Sam

bar

783.5

351.3

3.0

96

0.49

50.8

95.4

651.9

16.6

7241

2003

514.7

952.5

3.0

93.9

20.

4532.7

83.6

552.9

10.9

5125

2001

220.5

2

47

60.9

1.3

43.4

80.

1217.8

71.2

362.1

4.6

92

29

2000

Ch

ital

UC

LLC

L

Bio

mas

s den

sity

Wt

in

kg

Dg

CV

%D

gC

lust

er

size

En

cnt.

R

ate

95%

CL

D C

V

%D

NY

ear

Spec

ies

125.1

816.6

1.2

011.5

50.1

320.4

19.4

70.1

813.9

065

2003

186.2

025.6

2.5

18.2

20.2

836.4

111

.726.7

20.6

8130

2001

227.8

9

36.4

2.2

511.2

30.1

758.5

710.9

37.8

25.3

12000

Lan

gur

63.3

245.1

1.0

91.8

10.1

75.3

20.

7345.5

1.9

785

2003

29.2

141.6

0.7

21.6

20.1

2.1

80.3

842.1

0.9

144

2001

73.6

3

32

49.4

1.2

21.8

80.1

6.9

40.

7650.3

2.3

025

2000

Ch

inkar

a

30.2

252.1

0.3

74.0

50.0

44.6

20.

4957.8

1.5

120

2003

27.3

442.6

0.5

83.2

80.0

83.4

40.

5446.5

1.3

629

2001

NA

20

Da

tase

t to

o sm

all

to e

stim

ate

den

sity

62000

Wild

pig

56.7

42

20.2

2.2

11.2

20.1

74.1

91.

7320.6

2.7

082

2003

82.9

919

3.3

21.2

30.2

15.9

52.6

219.2

3.9

596

2001

62.4

75

21

29.5

1.8

41.6

10.1

35.7

11.

531.9

2.9

731

2000

4H

An

telo

pe

1639

24.5

4.4

02.9

80.6

823.0

97.

4426

13.1

1333

2003

787.6

237

2.6

72.7

20.6

13.9

92.8

337.2

6.3

0126

2001

1421.7

5

125

26

3.6

23.1

40.4

421.0

66.

1429.1

11.3

7107

2000

Nilg

ai

1387.5

733.7

3.9

52.6

20.4

522.0

04.

8734.1

10.3

5223

2003

1175.5

836.9

3.9

72.

30.4

719.4

13.9

637.2

8.7

7234

2001

2510.8

9

134

34.2

7.1

82.6

10.6

41.5

28.

4534.9

18.7

3147

2000

Sam

bar

783.5

351.3

3.0

96

0.49

50.8

95.4

651.9

16.6

7241

2003

514.7

952.5

3.0

93.9

20.

4532.7

83.6

552.9

10.9

5125

2001

220.5

2

47

60.9

1.3

43.4

80.

1217.8

71.2

362.1

4.6

92

29

2000

Ch

ital

UC

LLC

L

Bio

mas

s den

sity

Wt

in

kg

Dg

CV

%D

gC

lust

er

size

En

cnt.

R

ate

95%

CL

D C

V

%D

NY

ear

Spec

ies

Tabl

e 3.

Ani

mal

and

bio

mas

s de

nsity

(kg/

km2 )

est

imat

es o

f tig

er p

rey

in P

anna

Tig

er R

eser

ve. (

n= n

umbe

r of

det

ectio

n; D

= de

nsity

of

anim

als/

km2 ;

D c

v= c

oeff

icie

nt o

f va

riat

ion

dens

ity; 9

5% C

L=

95%

con

fide

nce

inte

rval

; Enc

nt. R

.= e

ncou

nter

rat

e; D

g= g

roup

den

sity

/km

2 ; D

g cv

= co

effi

cien

tof

var

iatio

n of

gro

up d

ensi

ty; w

t in

kg=

aver

age

wt o

f the

spe

cies

).

21

potentially limits the length of transects, whichwill largely depend on length, visibility, terrain andnumber of detection. To minimize the errorencountered in morning and evening counts inthe shorter, intensive 2001 surveys of fifteendays and the increase in group size and

0

0 . 2

0 . 4

0 . 6

0 . 8

1

1 . 2

1 . 4

1 2 3 4 5 6 7 81 5 m i n u t e s T i m e s l o t

M o r n i n g E v e n i n g

0

0 . 5

1

1.5

2

2 . 5

1 2 3 4 5 6 7 8

1 5 m i n u t e s T i m e s l o t


0

0 . 2

0 . 4

0 . 6

0 . 8

1

1.2

1.4

1.6

1.8

2

1 2 3 4 5 6 7 8



L i n e a r ( E v e n i n g ) L i n e a r ( M o r n i n g )

( b ) s a m b a r

( c ) n i l g a i

( a ) c h i t a l

0

0 . 2

0 . 4

0 . 6

0 . 8

1

1 . 2

1 . 4



0

0 . 5

1

1.5

2

2 . 5

1 2 3 4 5 6 7 8



0

0 . 2

0 . 4

0 . 6

0 . 8

1

1.2

1.4

1.6

1.8

2

1 2 3 4 5 6 7 8




0

0 . 2

0 . 4

0 . 6

0 . 8

1

1 . 2

1 . 4



0

0 . 5

1

1.5

2

2 . 5

1 2 3 4 5 6 7 8



0

0 . 2

0 . 4

0 . 6

0 . 8

1

1.2

1.4

1.6

1.8

2

1 2 3 4 5 6 7 8




( b ) s a m b a r

( c ) n i l g a i

( a ) c h i t a l

Figure 7. Encounter rates (animal groups (or sighting)/hour) pooled for every 15 minute time interval of three majorspecies of tiger during the walk on the line in the morning (between 6.45 and 8.30 am) and evening (between 16.30to 18.15 pm), in Panna Tiger Reserve. Straight dotted lines indicate trend for evening and continuous line is formorning trend. (a) chital; (b) sambar and (c) nilgai. (a) chital

encounter rate during the relatively extended2003 survey, a new survey design is needed. Asampling period (as short as possible) with morelines, which are walked once a day and repeatedafter three to four days, can minimize the errorsencountered during the surveys.

22

71.0246.6452.27Average

39.2421.2673.08Panna NP

69.344.5954.76Ranthambhore NP

73.9144.1356.13Nagarahole NP

70.9130.1469.85Kajiranga NP

91.0980.8118.76Kanha NP

81.6658.9141.08Pench NP

Medium preyLarge prey

Animal densityMedium prey

Biomass Density in PercentImportant tiger habitat

71.0246.6452.27Average

39.2421.2673.08Panna NP


73.9144.1356.13Nagarahole NP


91.0980.8118.76Kanha NP

81.6658.9141.08Pench NP

Medium preyLarge prey

Animal densityMedium prey

Biomass Density in PercentImportant tiger habitat

0

20

40

60

80

100

120

5 6 7 8 9 10 11 12 1 2 3 4 5 6

Hours morning to evening

Per

cent

act

ive

Figure 8. Percent activity of radio-collared sambar during February and March months in Panna Tiger Reserve.

Table 4. Biomass density and density of medium and large tiger prey in different Protected Areas ofthe Indian sub-continent, (figure in percent).

23

The home range of an animal is the areatraversed by an individual in its normal activitiesof food gathering, mating and caring for theyoung (Burt, 1943). Animals often rankresources, and home range can be envisioned asan integration of contour maps of theseresources, which include escape cover, travelroutes and known home ranges of members ofother sex (Ellner and Real, 1989; Pyke, 1984; Pykeet. al. 1977). If the sum of daily costs ofmaintaining, monitoring, defending, developingand remembering the critical resources exceedthe benefits gained from the home range it willreflect in the animals’ nomadic nature and it willnot exhibit site fidelity. Site fidelity is usually anindicator of whether an animal has establisheda home range (Powell, 2000). We have used Minimum Convex Polygonmethod which is the most commonly usedmethod of estimating home ranges (Jennrich andTurner, 1969). This method is sensitive toextreme data points as it incorporates areasthat are not used by the animal. The HarmonicMean Estimator is used to explain the patternof use or centre of activity based on theharmonic mean distances (Dixon and Chapman,1980). Harmonic Mean Estimator mayaccurately show multiple centres of activity, buteach estimated utility distribution is unique tothe position and spacing of the underlying grid. Animal ecology pivots around two importantresource concerns: forage and partner as mate.Food is a critical resource, important for allanimals. Females who have the addedresponsibilities of nurturing young ones areexpected to consider forage as a prime resource(Putman, 1980). Males, on the other hand carrythe responsibility of passing over their genes andmaking sure that they get the best opportunitiesto mate with females in oestrous. Since forageand oestrous mate are both resources withseasonal availability, movement patterns acrossdifferent seasons is expected to reflect theavailability of these resources. Seasonal andannual average of distance covered within 24hours provides a clear picture of resourceavailability and energy spent on these resourcesby different individuals.

Methods Five sambar and two chital individuals wereradio-collared for the purpose of this study.These comprised of two adult sambar males(Msam-004, Msam-127) and one adult chitalmale (Mch-05) and three females sambar

TIGER PREY-II: Home range and movement, habitat use andactivity pattern of sambar and chital

Tab

le 5

. Rad

io-t

rack

ing

of

five

rad

io-c

olla

red

sam

bar

in P

NP.

24

(Fsam-024, Fsam-026, Fsam-129) and onefemale chital (Fch-02). These animals weremonitored on a regular schedule in differentseasons. Radio-collared animals were radio-tracked for 14 to 17 months in the year 2001and 2002. Details of tracking effort are providedin table 5. No mortalities of any radio-collaredanimals are observed within the study period.Detailed description of techniques fordetermining locations of radio-collared animalsis given in the Chapter-III “methods”. We usedMinimum Convex Polygons (MCP, Jennrich andTurner, 1969) for evaluating the home ranges ofsambar and Harmonic Mean (HM) estimator(Dixon and Chapman, 1980) for determining thecentre of activity. In addition to calculation ofannual home ranges, seasonal home ranges arealso calculated for monsoon, winter and summerseasons. Animals make occasional forays into areasbeyond their regular home range. Locations ofthese forays lead to overestimation of homerange sizes. Since such forays do not representactual home ranges, such locations are excludedfrom the home range analyses. This problem isaddressed by calculating home ranges with 95%of the total radio-locations. The outer most 5%locations and observations on these locationsare described separately. For triangulation andcalculation of home ranges we used LOASTM andBIOTASTM software (Ecological SoftwareSolutionTM, © 1998-2001). The distancebetween two locations of successive days is usedto determine the distance covered by a radio-collared animal in 24 hours. These distancesare calculated using custom made macros in anMS EXCELTM spreadsheet.

Results The average annual home range size of malechital and sambar is larger (male sambar=13.185km2, n=2; male chital=XX) than that of females(female sambar= .46 km2, n=3; female chital=XX). Detailed results with seasonal and annualhome ranges (MCP) and centres of activity (HM)are given in table XX. Figure XX provides acomparable graph of annual and seasonal homeranges of all the five radio-collared sambar andtwo chital.

Home rangesMale-004 Sambar-004 was tracked for 14 months

between March 2001 and April 2002. Theaverage seasonal home range size estimated forsambar-004 is 9.01 km2, which is similar in sizeto its annual home range (11.49 km2, n= 151;figure 6.2). This sambar ranged over larger areasduring all seasons and its home range sizes insummer (9.08 km2, n=39), monsoon (10.02km2, n=41) and winter (7.77 km2, n=71) show onlymarginal variation (Figure 9). Two centres of activity are observed withinits annual range and the total area is estimatedat 4.99 km2 (80% HM, Figure 9). The smaller ofthe two centres of activity is located around itsrutting activities. It spent most of its time onthe upper Talgaon plateau, far away from therutting ground.

Male-127 Sambar-127 was tracked for 17 months.Unlike sambar-004, its average seasonal homerange size (6.13 km2) is less than half (41%) ofits annual range (14.88 km2, n=226, figure 6.4).

Monsoon (3.04 km2) and winter (2.32 km2)ranges are small. It ranged over much a largerarea during the summer months (13.03 km2,figure 8). Only one centre of activity (5.77 km2, 80%HM) is observed within its annual range (Figure8), which overlapped almost entirely with thehome range of its rutting season i.e. winter, andalso with its monsoon home range. Its centre ofactivity is similar in size to that of sambar-004.Female-024 Female-024 was radio-tracked for 17months. Its average seasonal home range sizeis small, only 43 % in size in comparison to itsannual home range (8.2 km2, n=198; figure 12).Winter (2.22 km2, n=65) and monsoon (0.44

7.7771Winter

9.0839Summer

10.0241Monsoon

4.9911.49151Annual

HMMCPnPeriod

Area (km2)Male-004

7.7771Winter

9.0839Summer

10.0241Monsoon

4.9911.49151Annual

HMMCPnPeriod

Area (km2)Male-004

25

km2, n=43) home ranges are much smaller thanthe summer home range (8.08 km2, n=90,Figure 12). Only one centre of activity is noticedwithin its home range, where 80% of its activitieswere restricted to this small area of 2.39 km2(80% HM; figure 12). This centre of activity alsooverlaps almost completely with its winter andmonsoon ranges.Female-026 Female-026 was radio-tracked for 14months. Like female-024, the average seasonalhome range size of female-026 is also 43 % ofthe annual home range (6.09 km2, n=194; figure13). Its summer home range is large (6.17 km2,

n=92) in comparison to the winter (1.08 km2,n=41) and monsoon (0.75 km2, n=61) homeranges (Figure 6.9). Its centre of activity isrestricted within 2.07 km2 (80% HM; Figure 13).The centre of activity overlaps closely with itswinter and monsoon ranges.

Female-129 Female-129 was monitored for 17 months.Unlike the other two radio collared females, itsaverage seasonal home range size is as large as(81%) of its annual home range (5.08 km2,n=240; Figure 11). Its home range in the monsoon

was very small (0.53 km2, n=56), but it rangedover a much larger area during winter (6.80 km2,n=84) and summer months (5.00 km2, n=100;Figure 11). The centre of activity is restricted to2.46 km2 (80% HM; figure 11) and encompassedthe monsoon home range completely.Movement Pattern

Average distance travelled by all individualssambar is 0.84 km (range: 0.008 km to 5.8 km).Males travelled longer distances (1.11 km, range= 0.011 km to 5.8 km) than females (0.67 km,range = 0.008 km to 4.8 km). Only Male-004moved longest distances (1.49 km, range: 0.18

km to 5.8 km) during monsoon while all otherradio-collared sambar travelled most duringsummer. Male-004 had the largest average ofannual distance travelled within 24 hours (1.22km, range = 0.073 km to 5.8 km) whereasfemale-024 travelled the least (0.63 km, range:0.008 km to 4.39 km). Detailed results ofseasonal and annual distance covered by theradio-collared sambar are given in table 6.

Habitat Use and Activity Patterns Management of wildlife populationsessentially incorporates management of habitat

2.2265Winter

8.0890Summer

0.4443Monsoon

2.398.20198Annual

HMMCPnPeriod

Area (km2)Female-024

2.2265Winter

8.0890Summer

0.4443Monsoon

2.398.20198Annual

HMMCPnPeriod


2.3275Winter

13.0385Summer

3.0466Monsoon

5.7714.88226Annual

HMMCPnPeriod

Area (km2)Male-127

2.3275Winter

13.0385Summer

3.0466Monsoon

5.7714.88226Annual

HMMCPnPeriod

Area (km2)Male-127

1.0841Winter

6.1792Summer

0.7561Monsoon

2.076.09194Annual

HMMCPnPeriod


1.0841Winter

6.1792Summer

0.7561Monsoon

2.076.09194Annual

HMMCPnPeriod


6.8084Winter

5.00100Summer

0.5356Monsoon

2.465.08240Annual

HMMCPnPeriod


6.8084Winter

5.00100Summer

0.5356Monsoon

2.465.08240Annual

HMMCPnPeriod


26

for wildlife populations. For proper habitatmanagement, it is mandatory to have someunderstanding of species’ requirements. Naturalselection favours those individuals who preferhabitats in which reproductive success is higherand where they survive better (Krebs, 1994).Preference can be defined as the likelihood of aresource being chosen often, if offered on anequal basis with others (Johnson, 1980).Preference for suitable habitat increases fitnessand reproductive success of the population andtherefore a high equilibrium of density ismaintained. Disproportionate use of a habitatcompared to its availability is considered as anindication of preference. There are only a handful ecological studieson sambar and chital that have been conducted

in the subcontinent (Schaller, 1967; Mishra,1982; Khan, 1990; Sanker, 1994). Managementof tiger is in fact directly related to managementof its prey and their biomass needs (Karanthand Stith, 1999). To evaluate the ecologicalrequirements of sambar and chital, it wasimportant to find out what habitat parametersdictate their well-being.

Analyses Locations of radio-collared sambar and chitalwere used for estimating habitat use andpreferences. These locations obtained fromradio-telemetry monitoring of the radio-collaredanimals were overlaid on the habitat map ofPanna Tiger Reserve. This habitat map wasdeveloped from satellite imagery. Seven distincthabitat types were classified using satelliteLISS-III images based on ground truthing plots(Chundawat, unpublished report). These sevenhabitat types are:

1. Disturbed (settlements): 8% of thetotal park area

2. Open/Grassland: 22% of the total parkarea

3. Thorn forest: 10% of the total park area4. Miscellaneous: 26% of the total park area5. Mixed forest: 10% of the total park area6. Mixed dense: 20% of the total park area7. Water body: 4% of the total park area

0.783.230.020.613.770.010.704.950.021.03Females

0.743.230.040.633.770.030.784.790.090.98F-129

0.982.950.040.693.260.010.914.950.181.30F-127

0.631.590.020.531.210.090.423.070.020.82F-026

0.935.800.031.015.200.010.764.390.20.98Males

0.631.590.030.540.710.010.344.390.060.83M-024

1.235.800.181.495.200.071.183.860.201.13M-004

Av. Dist.

RangeAv. Dist.

RangeAv. Dist.

RangeAv. Dist.

AnnualWinterMonsoonSummerSambar

0.783.230.020.613.770.010.704.950.021.03Females

0.743.230.040.633.770.030.784.790.090.98F-129

0.982.950.040.693.260.010.914.950.181.30F-127

0.631.590.020.531.210.090.423.070.020.82F-026

0.935.800.031.015.200.010.764.390.20.98Males

0.631.590.030.540.710.010.344.390.060.83M-024

1.235.800.181.495.200.071.183.860.201.13M-004

Av. Dist.

RangeAv. Dist.

RangeAv. Dist.

RangeAv. Dist.

AnnualWinterMonsoonSummerSambar

Sambar

04

Sambar

024

Sambar

026

Sambar

127

Sambar

129

0

2

4

6

8

10

12

14

16

area

(sq

km

)

monsoon

winter

summer

Annual

Table 6. Distances covered by radio-collared sambar within 4 hour period in PNP

Figure 8. Comparison of seasonal home ranges of sambar.

27

Figure 9. Home ranges of Male-004 sambar.

Figure 10. Home ranges of Male-17 sambar.

28

Figure 11. Home ranges of Female-129 sambar.


29

Water bodies were excluded from use andavailability analyses. I followed methodsdescribed by Neu et. al. (1974), later modifiedslightly by Byers and Steinhorst (1984) to testfor preference and avoidance and to determinewhich habitat types were preferred or avoided.For analyses, chi square was used to test forany significant preferences or avoidances withinthe six habitat types identified for the analysis.Bonferroni’s confidence intervals were estimatedto find out preference/avoidance of specifichabitat types. This was done by estimation ofBonferroni’s lower and upper confidence limits(LCL and UCL respectively) at 95% of proportionof use using the following equation:

CL= U ± [Z x ?U(1-U)/n]

We estimated the proportion of areascovered by the six habitat classes with respectto the total area of the park. Specificallydesigned macros using MS Excel were used forestimation of actual, expected, LCL and UCL ofhabitat proportions. Habitat preferences byradio-collared individuals were estimated using

males and females as separate group and allsambar as another group to determine seasonaland annual preferences.

ResultsHabitat preference and avoidance All radio-collared sambar showed significantpreferences (Figure 15) for mixed forests (41%)and mixed dense forest habitats (38.6%). Butavoided disturbed (0.1%), open/grassland (6.1%)and thorn forest habitats (0.2%) throughout theyear. Habitat use for male and females iscalculated separately to find out if any differentpatterns exist. Our results do not show anydifferences but point to a similar pattern. Acloser look at the individual level also did notreveal any different pattern in habitat use.

Annual We also estimated separately the annualhabitat preferences of all the five radio-collaredsambars (Figures 15). It is observed that male-004 used miscellaneous habitat in proportionto its availability but other radio-collared sambarshow significant preferences for miscellaneous,


30

mixed forest and mixed dense forest habitats;they also showed avoidance for disturbed, open/grassland and thorn forest habitats. Annual habitat selection by sambar malesand females was similar: they preferred mixeddense (male 35.69%; female 47.02%), mixed(male 36.31%; female 40.94%) and miscellaneousforest (male 16.88%; female 6.2%) and utilised

these habitats significantly more than theiravailability. Disturbed, open/grassland and thornforest habitats were avoided: as these habitatswere used less than their availability (table 7).

Seasonal The seasonal habitat preferences wereevaluated for pooled locations of males, femalesand for all combined. Only during the winters andmonsoon, did male sambar use densemiscellaneous habitat in proportion to itsavailability. Similarly, miscellaneous habitat wasused in proportion to its availability by males inmonsoon. Preferences in all other seasons had asimilar pattern as above (Figure 7.4 to 7.6 andtable 8).

Male sambar (M-004 and M-127) useddense mixed habitat in proportion to itsavailability during monsoon (27.28%) andsummer (46.9%), whereas the miscellaneousforest was used in proportion to its availabilityonly in the monsoon season (22.02%). Mixedforest was preferred and was used more thanits availability in all the three seasons. Preferencefor dense mixed forests was not as strong duringmonsoon for both male and female as wasobserved for other seasons (Figure XX). Femalesavoided the miscellaneous forest by utilizing itless than its availability in all three seasons.

Activity Pattern Seasonal variation in activity pattern isobserved in radio-collared sambar. Males andfemales showed similar activity patterns in

723457266168124165159107.0n=

0.2-0.3---0.6-Water body

38.640.936.339.943.539.437.135.5Mixed Dense

41.547.335.749.445.247.339.631.8Mixed Forest

11.66.316.98.95.64.2%13.220.6Miscellaneous

0.2-0.3---0.6-Thorn Forest

6.12.310.01.25.6-8.811.2Open/Grassland

0.10.2-0.6----Disturbed

1.73.00.5--9.1-0.9Outside

allFemaleMaleF129F026F024M127M004Habitat

723457266168124165159107.0n=

0.2-0.3---0.6-Water body

38.640.936.339.943.539.437.135.5Mixed Dense

41.547.335.749.445.247.339.631.8Mixed Forest

11.66.316.98.95.64.2%13.220.6Miscellaneous

0.2-0.3---0.6-Thorn Forest

6.12.310.01.25.6-8.811.2Open/Grassland

0.10.2-0.6----Disturbed

1.73.00.5--9.1-0.9Outside

allFemaleMaleF129F026F024M127M004Habitat

Table 7. Habitat use by individual radio-collared sambar in PNP.

Radio-collar sambar in PNP

31

male-127

-0.1

0

0.1

0.2

0.3

0.4

0.5

0.6

Dis

turb

ed

Ope

n/G

rass

land

Thor

n Fo

rest

Mis

cella

neou

s

Mix

ed F

ores

t

Mix

ed D

ense

used

LCL

UCL

expected

male-004

0

0.1

0.2

0.3

0.4

0.5

0.6

Dis

turb

ed

Ope

n/G

rass

land

Thor

n Fo

rest

Mis

cella

neou

s

Mix

ed F

ores

t

Mix

ed D

ense

used

LCL

UCL

expected

male-127

-0.1

0

0.1

0.2

0.3

0.4

0.5

0.6

Dis

turb

ed

Ope

n/G

rass

land

Thor

n Fo

rest

Mis

cella

neou

s

Mix

ed F

ores

t

Mix

ed D

ense

used

LCL

UCL

expected

male-004

0

0.1

0.2

0.3

0.4

0.5

0.6

Dis

turb

ed

Ope

n/G

rass

land

Thor

n Fo

rest

Mis

cella

neou

s

Mix

ed F

ores

t

Mix

ed D

ense

used

LCL

UCL

expected

46.746.646.935.240.729.730.433.427.3Dense Mixed

32.838.024.948.651.545.749.257.740.4Mixed Forest

12.211.213.710.02.017.912.22.722.0Miscellaneous

0.30.00.90.00.00.00.00.00.0Thorn Forest

7.73.713.66.35.86.78.26.210.3Open/Grassland

0.30.50.00.00.00.00.00.00.0Disturbed

AllFemaleMaleAllFemaleMaleAllFemaleMale

SummerWinterMonsoonHabitat

46.746.646.935.240.729.730.433.427.3Dense Mixed

32.838.024.948.651.545.749.257.740.4Mixed Forest

12.211.213.710.02.017.912.22.722.0Miscellaneous

0.30.00.90.00.00.00.00.00.0Thorn Forest

7.73.713.66.35.86.78.26.210.3Open/Grassland

0.30.50.00.00.00.00.00.00.0Disturbed

AllFemaleMaleAllFemaleMaleAllFemaleMale

SummerWinterMonsoonHabitat

Table 8. Habitat use by male and female sambar and all radio-collared sambar in PNP.

Figure 14. Habitat use (annual) by radio-collared male sambar in PNP.

32

female-129

-0.1

0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

Distu

rbed

Ope

n/G

rass

land

Tho

rn F

ores

t

Misce

llane

ous

Mixed

For

est

Mixe

d D

ense

used

LCL

UCL

expected

female-024

0

0.1

0.2

0.3

0.4

0.5

0.6

Distu

rbed

Ope

n/G

rass

land

Tho

rn F

ores

t

Misc

ellane

ous

Mixed

For

est

Mixe

d Den

se

used

LCL

UCL

expected

female-026

0

0.1

0.2

0.3

0.4

0.5

0.6

Distur

bed

Ope

n/G

rass

land

Tho

rn F

ores

t

Misc

ellane

ous

Mixe

d For

est

Mixe

d D

ense

used

LCL

UCL

expected

Fig. 14a. Annual habitat preference by female radio-collared sambar in PannaTiger Reserve.

33monsoon and summer. In winter, male sambar

are found to be more active throughout the 24hour period (>60% active). In comparison,females show reduced activities (<60% active)during night hours (2000 and 0700 hours)reaching to a minimum (<20%) in the earlymorning hours, around 0400 hours. Summeractivities show a very clear pattern with twodistinct activities peaks (>60% activities)between 0500-1100 hours and 1600-2200hours (Figure 17). In the middle of the day bothsexes show least activities (<30%) 1200-1400hours. No clear pattern is observed during themonsoon season and most of the time over 60%activity is recorded. Females remained marginallyless active (<60%) during night hours (Figure16)

DiscussionHome range and movement In Panna, as observed in many other areasand in other deer populations, the home rangesof male sambar are larger than those observedfor females (Putman, 1988; Johnson, 1984;Mishra, 1982; Sanker, 1994). There are not manystudies conducted in the Indian subcontinent tomake such comparison except one conducted inSariska (Sanker, 1994). Both studies documentsimilar differences between male and femaleranges. But the seasonal summer ranges arefound to be larger in Panna. Animals do not move randomly in the wild,they all have home areas in which they range tofulfil their minimum ecological and energeticrequirements. In this area of regular use definedas home range (Sanderson, 1966), seasonalvariations in environmental conditions alter theavailability of forage and other ecologicalresources such as water and cover and canaffect animals’ reproductive activities. Theseecological factors govern the movement patternand home range sizes of many of the deer species(Putman, 1988).

In Panna Tiger Reserve, the seasonalvariation that is observed in sambar homeranges is probably a response to the changingenvironmental conditions. Most of them use avery small area of a few hundred hectares duringthe monsoon (rainy season) months. It is duringthis season that the home range of sambar issmallest (Table XX). The monsoon is a season ofplenty in Panna Tiger Reserve. Most of the

All

-0.1

0

0.1

0.2

0.3

0.4

0.5

Dist

urbe

d

Ope

n/G

rass

land

Thor

n Fo

rest

Mis

cella

neou

s

Mix

ed F

ores

t

Mix

ed D

ense

used

LCL

UCL

expected

Male

-0.050

0.050.1

0.150.2

0.250.3

0.350.4

0.450.5

Dist

urbe

d

Ope

n/G

rass

land

Thor

n Fo

rest

Mis

cella

neou

s

Mix

ed F

ores

t

Mix

ed D

ense

used

LCL

UCL

expected

Female

-0.1

0

0.1

0.2

0.3

0.4

0.5

0.6

Dist

urbe

d

Ope

n/G

rass

land

Thor

n Fo

rest

Mis

cella

neou

s

Mix

ed F

ores

t

Mix

ed D

ense

used

LCL

UCL

expected

All

-0.1

0

0.1

0.2

0.3

0.4

0.5

Dist

urbe

d

Ope

n/G

rass

land

Thor

n Fo

rest

Mis

cella

neou

s

Mix

ed F

ores

t

Mix

ed D

ense

used

LCL

UCL

expected

Male

-0.050

0.050.1

0.150.2

0.250.3

0.350.4

0.450.5

Dist

urbe

d

Ope

n/G

rass

land

Thor

n Fo

rest

Mis

cella

neou

s

Mix

ed F

ores

t

Mix

ed D

ense

used

LCL

UCL

expected

Female

-0.1

0

0.1

0.2

0.3

0.4

0.5

0.6

Dist

urbe

d

Ope

n/G

rass

land

Thor

n Fo

rest

Mis

cella

neou

s

Mix

ed F

ores

t

Mix

ed D

ense

used

LCL

UCL

expected

Figure 15. Habita use all samabrs (a )conbined(all), (b) all males and (c) all females.

34

Monsoon: all sambar

0

0.1

0.2

0.3

0.4

0.5

0.6

0.7

Distu

rbed

Ope

n/G

rass

land

Tho

rn F

ores

t

Misce

llane

ous

Mixed

For

est

Den

se M

ixed

used

LCL

UCL

Expected

Winter: all sambar

0

0.1

0.2

0.3

0.4

0.5

0.6

Distu

rbed

Ope

n/G

rass

land

Tho

rn F

ores

t

Misc

ellane

ous

Mixed

For

est

Den

se M

ixed

used

LCL

UCL

Expected

Summer: all sambar

-0.1

0

0.1

0.2

0.3

0.4

0.5

0.6

Disturb

ed

Ope

n/G

rass

land

Tho

rn F

ores

t

Misc

ellane

ous

Mixed

For

est

Den

se M

ixed

used

LCL

UCL

Expected

Figure 15 a. Seasonal habitat preference by all radio-collared sambar inPanna Tiger Reserve.

35

Winter

0%20%40%60%80%

100%

1 3 5 7 9 11 13 15 17 19 21 23

Summer

0%

50%

100%

150%

1 3 5 7 9 11 13 15 17 19 21 23

Monsoon

0%20%40%60%80%

100%

1 3 5 7 9 11 13 15 17 19 21 23

winter

0%

2 0 %

4 0 %

6 0 %

8 0 %

100%

120%

1 3 5 7 9 11 13 15 17 19 21 2 3

summer

0%

2 0 %

4 0 %

6 0 %

8 0 %

100%

120%

1 3 5 7 9 11 13 15 17 19 21 23

monsoon

0%

2 0 %

4 0 %

6 0 %

8 0 %

100%

120%

1 3 5 7 9 11 13 15 17 19 21 2 3

Figure 16. Activity chart for male radio-collaredsambar.

Figure 17. Activity chart for male radio-collaredsambar.

36

important resources that govern the movementof deer (Putman, 1988) are available inabundance and distributed uniformly throughoutthe landscape used by the radio-collared sambar.Animals do not require to move larger distancesand can fulfil their minimum energetic andecological requirements within a relatively smallarea. Moreover, monsoon is also the season whenfawns are born, therefore parental cares demandfemale sambar to remain close to its fawningsite for an extended period. In addition toabundant food supply and its wider distribution,the fawning activity also affects the averagedistance travelled and movement pattern ofsambar, especially of females. It is most likelythis that is reflected in the smaller home rangesizes during the monsoon season as determinedby the radio-tracking of these animals. The onlystudy conducted on sambar in India forreasonable comparison is the Sariska one whereSanker (1994) did observe similar sized homeranges during the monsoon. Winter, a season, which follows the monsoon,still has a fairly abundant and wide resourcedistribution. This is also reflected in relativelysmall home ranges and similar trends that wereobserved in the monsoon. It is observed thatmales of many deer species move frequentlyduring the rutting season and wander over largerareas in search of receptive females (Putman,1988, Sanker, 1994). In Panna, the homes rangesof male sambar did not change much duringwinter months, which is the sambar ruttingseason, except for a minor increase. Averagedistances travelled in 24 hours only showmarginal increase in activity over those travelledduring the monsoon. The largest home ranges and averagedistance travelled by radio-collared sambar wasobserved during the summer months. In contrastto monsoon, summer is a season characterizedby poor quality and availability of forage, sparsedistribution of water, limited cover due to thedeciduous nature of the forest and of extremelyhigh ambient temperatures during day hours.Most of these resources are not distributeduniformly and frequent forest fires add to thenatural scarcity of these important ecologicalresources. Animals, therefore, are required tomove larger distances in order to find sufficient

food, water and cover to escape from theextreme temperatures. This behaviouraladaptation may be what manifests in largeraverage distances travelled daily and biggerranges as documented in this study throughintensive radio-tracking of collared sambar. It is observed that the females’ home rangesoverlapped considerably. Analysis of habitat useindicates that it is probably the availability ofresources, including forage and cover, that resultin higher site fidelity. Seasonal variations in homerange sizes may have been dependant uponmating behaviour and availability of water, shelterand other resources in addition to the variableclimatic conditions.

Habitat use Sambar is classified as a browser (Putman,1988) and has evolved successfully in forestedhabitats (Sunquist et al., 1999). Their anti-predatory strategies and food requirementsnecessarily demand forest cover for survival. Itis found that sambar is generally very selectivein their choice of habitat (Mishra, 1982; Sanker,1994). Results of habitat preference usingBonferroni’s confidence interval suggest thatmixed forests, dense mixed forests andmiscellaneous forests, which are associated withhigher canopy cover and tree densities, arepreferred by sambar in Panna Tiger Reserve.These results also reveal that sambar avoid openforests, grasslands, thorn forests: they aresensitive to anthropogenic pressures anddisturbances and avoid such habitats. Resultsof this study corroborate that sambar and nilgaiuse distinctly different habitats (Mathai, 1999;Chundawat, 2001). This difference can beaccredited to their anti-predatory behaviour.Nilgai prefers open terrain and large groups formore vigil; sambar rely upon small groups,camouflage and thickets for hiding (Putman,1988). Distribution of sambar is thereforerestricted mainly to dense forested habitatswhere they achieve higher abundance. Thesepreferred habitats of sambar are also frequentlyused habitats for tiger and therefore theseforests habitats are of significant importancefor the management of tiger and its associatedprey in the Reserve. The most regularly preferredhabitat, i.e. mixed forest is only 10.1% of the total

37

park area. To manage a healthy tiger population,which largely depends on the sambar in PannaTiger Reserve, it is important to maintain ahealthy mosaic of dense forested habitats. No difference in habitat preference acrossseasons is observed in this study. Though, malesambar used dense mixed and miscellaneousforest within their proportion of availabilityduring monsoon and winter. This proportionalusage in winter can be credited to the large areathat males cover during these two seasonspossibly in search of mates (Putman, 1988).Females also show similar patterns of usagesince they are more dependent on optimal forageavailability (Putman, 1988; Mishra, 1982). It cansafely be deduced here that forage is the criticalresource for female sambar, whereas partner formating mainly governs the terrain utilization bymales. Seasonal variation in activity patterns ofradio-collared sambar gives a clear idea aboutenergy budgeting. The day and nighttemperatures in monsoon do not vary much andit is reflected in overall activity pattern whichdid not vary across the entire 24 hour period.Whereas, in winter, day and night temperaturesvary considerably but remained below 25oC, whichallowed the animals to remain active throughout.Moreover, winter is the rutting season whenmales move great distances in search of mates(Putman, 1988; Mishra, 1982, Sanker, 1994).This may be one of the prime reasons forextended active periods as observed in the study.This is especially true for males which remain>60% active for most of the 24 hour. In contrastto males’ activity pattern, females show reducedactivities (<40% activity) for most of the nighthours. Temperature variations between minimum

and maximum is at its highest (over 20oC)during the summer. In Panna, summers areextremely hot and temperatures can go up to480C during mid day; hence there is a clearreduction in activity around mid day. Most ofthe activities are seen during early morning andin the evenings, when temperatures are optimal.These results show that there is diagnostictemporal energy budgeting during summers,where resources are scarce and sporadicallydistributed. During winter and monsoon,abundance of resources and independence fromthe temperature regime results in a lessdemarcated temporal variation in activitypattern within a 24-hour day. This is the firststudy to document the activity pattern ofsambar. Availability and dispersion of resources is acritical parameter that determines pattern ofuse by an animal. For widely dispersed availabilityof resources, the average daily movement of ananimal would be larger. Large average movementwould necessarily mean higher energy costs. Thusit is important that the benefits from theseenergy expensive movements be higher than thecosts (Putman, 1988; Powell, 2000). From thestudy it appears that summer which is a seasonwhen important resources are scarce, thisaffects the space requirement for individualsambar and possibly controls the number ofanimals per unit area or abundance of sambarpopulation. Hence the summer season seeminglydetermines the population size of sambar, oneof the major prey of tigers in Panna Tiger Reserve.It is therefore reasonable to presume thatavailability of key resources during summer playa very important role in ecology of tiger inaddition to other biotic factors such as grazingand anthropogenic disturbance.

38

Carnivores show a great diversity in theirsocial organization and behaviour, some aresolitary and others live in highly developed socialunits. The social organization and behaviour ofa carnivore reflects its response toenvironmental and ecological factors and to itscon-specifics.

Dispersal Dispersal is defined as a movement awayfrom the natal area to a place where dispersingindividuals settle for breeding. Two kinds ofmovement are recognized: one in which an animalfinds a suitable breeding spot near its natal area? usually known as phillopatry; the other is a long-range movement away from natal area. Thedistance travelled by the dispersing individualdetermines the gene flow within or withneighbouring populations and determines thedemography and social organization of thepopulation. In this study information on dispersal wascollected through radio-collaring dispersingtigers and identifying individual tigers by facialmarkings and stripe patterns. Since dispersalwas not the main focus of the study informationon the subject was limited to radio-taggedindividuals and identified tigers only. For thisstudy, movement away from the mother’sterritory was taken as dispersal; if the animalwas localized in an area to establish a territoryand started breeding subsequently, this wasconsidered phillopatric movement. Where anindividual moved away, out of the study area,for a long time, this was considered long distancedispersal.

ResultsPopulation characteristics Most of the information obtained on tigerbiology in this study comes from the radio-collared tigers and from several individuallyidentified tigers (based on repeatedphotographs). Female-118 was the first adultfemale tiger radio-collared and the three femalecubs (the three siblings Fm-111; Fm-120; Fm-113)of her first litter were radio-collared when theywere either establishing their own territories or

had established it, in areas adjoining their natalarea (figure XX). During the first year of the study, in 1996, atotal of six tigers of over one year of age wereidentified within the intensive study area of 180km2. This included two territorial females andtwo males. Both the females were raising cubs,the female “Female-118”, who occupied the besthabitat in the Reserve in Chapner and Bhadarregion, was raising a litter of three cubs born inApril/May 1996; she was radio-collared inJanuary 1997. The other female “Bargadi female”(an identified individual) occupied the Bargadiregion and was raising a litter of two cubs. Thedominant large male, “Male-91” (radio-collaredin April 1996) patrolled both the femaleterritories and beyond, but no evidence ofanother resident female was found within histerritory during the intensive monitoring andradio-tracking of his movements. A very old malehad occupied the area south of Male-91’sterritory. The research team did not find evidenceof other non-resident tiger in this intensive studyarea. Thus during the first year of the study fouradult tigers were using an area of 180km2. Thisgave an approximate density of 2-3 adult tigersfor every 100 km2. Extrapolating from this figure,it was estimated that there were maybe 18-20adult tigers in the entire Reserve. The first litter of Female-118 comprised ofthree female cubs. They all survived to adulthoodand dispersed from the natal area andestablished their territories adjoining themother’s. Both the cubs of the Bargadi female,which comprised of one male and a female, laterdied at the early age of fifteen or sixteen monthsand twenty two months respectively. Bargadi, anold tigress, who had already lost a canine wasdisplaced by the female dispersing cubs of Fm-118. In mid 1998, male-91 had a fight with a newmale who took over part of it range near the river.Thus by the end of 1998, there were four femaleterritories within the same study area and twodifferent dominant males patrolled the region,increasing the density to over 3-4 tiger per 100km2. Since 1996, the number of breeding femalesin the areas doubled from two to four. In the timebetween 1996 and 2002 and Fm-118 gave birth

TIGER-I: Social Organization and Estimation of Tiger Population.

39

to four litters and her daughters females-111, 113,and 120 each had two to three litters. As aresult, the number of transient and sub-adultanimals increased considerably. In the studyarea, by early 2002, there were seven residentbreeding females, two territorial males and fourto five young known tigers. In 2002 the sevenfemale tigers each had a litter; this includedfemale-118 with three cubs and the otherfemales-111, 113 and 120, the Amdar, Gata andNararan females each had two cubs: altogetherthis accounted for 15 cubs. This meant thatthere were nine resident tigers ? two territorialmales and seven territorial females ? and fouror five known sub-adult tigers, plus fifteen cubsof various ages. These all accounted for a totalpopulation for over 28 tigers in an area of over400 km2 intensively monitored. Between 1996 and 2004, a total of 42different tigers were identified and monitored,which included adult resident breeding tigers,young adult non territorial, non breeding animalsand cubs over one year of age. Thirty-eight ofthem were positively identified, photographed andsexed. These included nineteen females andtwenty males, giving almost an equal sex ratio.This may be due to the fact that during the laterpart of the study, between 2002 and 2004,there was a frequent change-over of fourdifferent dominant males. ‘Hairy foot’ took overafter the dominant male 091 disappeared in2002, but shortly after he was found dead, inearly monsoon of 2003. Another young male,identified as “Broken-tooth”, replaced him andtook over the area. Within the two dominant male territoriesbeing monitored, seven breeding femaleterritories were identified by the end of 2001.Out of these seven females, four of the femalesand both the males were radio-collared. The ratioof breeding individuals favoured females, 3 : 1. Itis observed that the breeding population in Pannais limited due to availability of space and due tothe poor quality of tiger habitat available.The rest of the tiger population was composedmainly of pre dispersal adult cubs and a fewtransient individuals. The transient populationwas mainly young adults from previous littersand it was the male sex that remainedassociated with natal areas longer and these

were observed intermittently.

Reproduction During the study we observed residenttigresses were either pregnant or raising a litterat any time of their reproductive age (> 3 to 14years). This makes tiger very productive andresilient species. During this period, we monitoredeight breeding tigresses giving birth to fourteenlitters and a total of 32 cubs. It was alsoobserved that female cubs tend to wean toindependence and leave the natal area at anearlier age than their male siblings.

Age of sexual maturity: In Panna we observed that, young femalesgave birth to their first litter, on average, at theage of 39.75 months (n=5); the earliestobserved during the study was at the 32ndmonth. We were able to observe four female cubsdispersing and later breeding by placing radio-collars on three of them. On average a fresh litteris produced at an average gap of 21.6 months(n=14). Almost all the litters observed wereproduced on 22nd months, except one, which wasproduced in 20th months. Details of the littersproduced is given in table ?? A female is capable of giving birth to a freshlitter in just a few months interval. This wasobserved in the case of tigress Fm-120, whenher entire second litter, which was only fourmonths old, was killed by a new dominant male,‘Hairy-foot’. These cubs of the second litter werelast seen on 26th December 2001. Her thirdlitter of two cubs were later seen on 10th March,when they about a week old, this meant thatshe conceived before her previous litter died.Signs of the new male ‘Hairy-foot’ visiting Fm-120 were seen on several occasions and once hewas seen resting for hours just a few yards awayfrom the mother and her litter.

Litter size and gestation period Average size of litters observed during thestudy is 2.3 cubs per litter (n=14). This is basedon the observation of cubs survived over 3-4months. It is extremely difficult to find youngcubs and only on three occasions were we ableto observe litters a few weeks old and we did notobserve any mortality at the age of 2 to 12 weeks.

40

Based on the twelve birth and matingsobserved by the research team, the averagegestation period estimated in Panna tigers was103 days. Mating was observed throughout theyear, thrice in December, twice in August and onceeach in January, February, May, June, July,September and October. These field observationson the radio-collard tigresses suggest thatestrous cycle and mating time depend on thefirst litter and subsequent mating shiftaccordingly by two to three months every birth.We did not see any seasonality in mating thoughit appears that most of the mating took placeduring monsoon and winter but we saw matingthroughout the year.

Mortality and SurvivalA total of 42 tigers were identified and

monitored regularly during the study period. Thisincludes thirteen adult, breeding tigers ? ninefemales and four males ? and 32 cubs. The cubpopulation also includes the three female cubswho grew to adulthood and were later monitoredas independent breeding tigresses (Female-111;113; 120). Details of the 42 tigers survivalobservation is given in Appendix-II. A total of nine mortalities in adult tigersare suspected but the research team could findonly two carcasses of tigers. These ninesuspected mortalities include the Bargadifemale, who was the dominant territorial femaleof the Bargadi region who was displaced byFemale-120 in 1998; it is suspected that shedied most likely due to old age after she wasdisplaced from her territory. Female-111, one ofthe siblings of Female-120 and 113, went missingover a cattle kill in October 2002 close to theperiphery of the Park boundary near Talgaon. Herpresence could not be confirmed by intensivecamera trapping in the areas she usually visited.After three months of intensive monitoring andlooking for other signs, this failure to confirmher presence led the research team to suspectmortality. A couple of months later, her sibling,Female-120 (radio-collared in 1999) was founddead in a snare, in December 2002. Her bodywas recovered and at the site, carcasses ofsambar deer that had also been snared wererecovered from the site. ‘Hairy foot’, a new malewho replaced the dominant radio-collared male

Male-091 (monitored since 1995 to 2002), wasfound dead in a well in June 2003. This wasfollowed by the disappearance of the third sibling,Female- -113 (radio collared in 2000) by March2004. Later, within a year, another male ‘Broken-tooth” was seen with an injury on its shoulder.All the above-mentioned tigers were intensivelymonitored animals and there is no report of theirsurvival during the period monitoring whichcontinued up to March 2005. (Broken-tooth wasseen later.) Of the other tigers that weremonitored, though not as intensively, there isno further evidence of their presence in theirareas; this includes the three females fromAmdar, Nararan and Gata. In addition to observations made on thesurvival and mortalities on adult tigers, we alsoobserved the birth of 32 cubs from 14 littersbetween January 1996 and December 2005 andobserved only eight mortalities up to two yearsof age. Both the cubs of the first observed litterof Bargadi females died (Table XX), one due tounidentified disease (carcass recovered) and theother died over a cattle kill (poaching suspectedbased on the circumstantial evidences). Theother two mortalities were due to suspectedinfanticide of the second litter of Female-120 bythe new male ‘Hairy foot’ after he replaced theradio-collared male-91. Other mortalities wereeither due to natural or due to unknown causes.Only on three occasions we were able to monitorlitters from a very early age of few weeks and wedid not see any mortality in these litters (sevencubs) of less then two months.

Survival/Mortality RateFor calculating mortality and survival rate

the study period was divided in two periods, 1996to 2001 (Almost six years or 5.9 years) and2002 to 2005 (3.3 years or almost threeyears). This was done because mortality rateincreased significantly from 2002. We usedcomputer software MICROMORT to calculatesurvival mortality rate (Ref XX) It is calculated that for a tiger, irrespectiveof its sex, to survive through the nine years inPanna is very poor, just 21.7% (95% confidenceinterval of 0.075 and 0.626; Var = 0.013752).But for the first six years of the study survivalprobability is very high 81.6% (Confidence interval

41

of 0.548 and 1.0). It drops significantly for nextthree years to astoundingly low 26%. Probability that a female will survive duringthe first six years of the study is 70% but forthe next three years it is only 24% and for theentire nine years it is only 16%. Probability thatmales will survive for the first six years is almostone. This is an overestimation due low samplesize and absence of mortality in the monitoredpopulation. But after 2001, the chances of malessurvival for the next three years declines to 28%which is similar to the survival in the entire nine-year monitoring period.

Dispersal We witnessed four female and one male cubsgrowing and dispersing and three of them,Female-111, Female-120 and Female-113, (laterradio-collared) were siblings and the first litterof Female-118. Female-111 separated from theother siblings earlier, at the age of about 18months and moved to a new area in the east;but for about six months, she continued to useand make kills within her natal area. Later shemoved further east and established herindependent territory adjoining her mother’s. Theother two siblings moved into an area occupiedby the old Bargadi female west of their natalarea. These two siblings moved in this area incoalition and displaced the old female. Theyremained in coalition for several months and wereseen occasionally in association with the oldfemale’s male cub. The female-120 establishedher territory and patrolled this area until herdeath. Her coalition partner and the third siblingmoved north to an area along the river Ken valleyand established her territory immediatelyadjoining her mother,s. These three siblings wereradio-tracked from 1998 to 2002. The fourthdispersal observed was that of another femalecub of female-118, which belonged to her fourthlitter. She had partially occupied space leftvacant after the death of female-120. A Malecub of female-118 was seen as a transient animaland was observed mating with female-111 and alsofemale-113.

Estimation of tiger population As part of the ongoing investigation on tigerecology, the research team also undertook

fieldwork to determine tiger density in PannaTiger Reserve. The tiger’s secretive nature andinherent and relatively low density make hard itto estimate numbers reliably. But recenttechnological and theoretical advances, and theirinnovative application by field biologists nowmake it possible to estimate tigers number morereliably. The stripe pattern of every tiger is uniqueand this allows us to identify individual tigerswhen remotely photographed. The record of itscapture history makes it possible to applycapture-recapture methods to estimate thepopulation size. The photographic capture-recapture sampling method, developed byKaranth and Nichols, 1998 to estimate tigerdensity, is being used in Panna Tiger Reserve. Thismethod has been used successfully in severaltropical forest habitat of the Indian sub-continent and in other south Asian tigerhabitats (Nichols and Karanth, 2002; Kawanishi,2002; O’Brien, et. al; 2003). The tiger population was estimated using thephotographic capture-recapture method. Thefield exercise was conducted in association withthe Centre For Wildlife Studies, who provided thecamera equipment for the census operation in2001. The data from the 2001 census ispublished in joint authorship with collaboratingscientist in Animal Conservation. We used camera traps manufactured byTrailmaster™. Each camera trap unit includesan electronic tripping device that uses an infrared beam to detect animal movements and firestwo separate camera units when it crosses thebeam. The cameras were placed on either side ofthe road at 3.5 to 4 metres apart so that bothflanks of the passing animal can bephotographed. These camera trap units consistof a transmitter and a receiver and two cameraswere deployed inside a steel shell to prevent theftplaced on both sides of the tiger’s expectedtravel path in order to photograph both itsflanks. The intensive study area was selected forsampling; this included the territories of all theradio-collared tigers. This is also the area fromwhere prey is estimated as part of the tigerResearch Project every year. In this area, 60camera trap sites were identified. Camera trapswere placed at spaces determined after an

42

intensive survey of the region and othermethodological considerations. Each cameratrap consists of cameras and an infra-red triggerunit, which is activated by an animal movementbetween the units. Two cameras are used at eachsite and are placed on either side of the trail tophotograph both flanks of the tiger (repetition– delete here?). Since we only had 20 camera traps units andneeded to cover 60 trap-sites, for logisticalreasons the sampling area was divided into threeunits, each with 20 trap locations. Each areawas sampled for tiger ID pictures for 15 daysand then the camera trap units were moved anddeployed in the next sampling block. All the unitswere visited every day to record the capturedetails in a standard format to document allthe necessary information needed for lateranalysis and to troubleshoot where necessary(Karanth & Nichols, 2002: page 183). Each roll of film was marked withidentification numbers and the unique numberof the frames for each photograph recorded. Thishelped us to assign each capture correctly andmatch both profiles. In addition to this each tigeridentified was given an identification number.Tigers were identified from their unique stripepatterns (Karanth et al., 2002). We used CAPTURE software (Rexstad &Burnham 1991) to analyse the data. From theabove field notes capture histories for theidentified tigers was constructed following thestandard protocol described in Otis et al., 1978and Nichols, 1992. ‘1’ indicated capture and ‘0’denoted no captured for that that occasion.Details capture histories are given in the Table-XX. For the analysis of this data we presumedthat that the sampled population isdemographically closed for the sampled period(Otis et al., 1978; Karanth, 1995; Karanth &Nichols, 1998) mainly because tigers arerelatively long lived animals. We considered the following models (Otis etal., 1978; Nichols, 1992) and estimation optionsimplemented in CAPTURE:

Mo = Capture probability is the same for alltigers, and is not influenced by behaviouralresponse, time, or individual heterogeneity.

Mh = Capture probabilities are heterogeneousfor each individual tiger, but not affected by trapresponse or time.

Mb = Capture probabilities differ betweenpreviously caught and uncaught tigers due totrap-response behaviour, but are not influencedby heterogeneity or time.

Mt = Capture probability is the same for allindividual tigers, but varies during the survey onlydue to time-specific factors. The model selection process also consideredmore complex models such as Mbh, Mth, Mtband Mtbh that incorporate effects ofheterogeneity, trap response and time, in variouscombinations. The overall model selection test(Otis et al., 1978) scores potential modelsbetween 0.0-1.0, with a higher score indicatinga better relative fit of the model to the specificcapture history data generated by the survey.For each analysis program CAPTURE generatedthe number of individual tigers captured (Mt+1),estimated capture probabilities per sample ,andthe estimated tiger population size (the numberof tigers in the sampled area, inclusive of animalsthat were not photo-captured at all). We used average half range length todetermine the buffer distance. This bufferdistance was used to generate a buffer aroundthe trap polygon to calculate the sampled area;(Wilson & Anderson, 1985; Karanth & Nichols,1998; Nichols & Karanth, 2002)

Results In February-April 2002, camera trapping wasdone for 45 days and include three 15-daysampling. This accounts for a total 914 trap-nights. Thirty-six usable photographs (17 rightflanks, 19 left flanks) of 11 individual tigers (8females, 3 males) were obtained from thissampling. As generally expected for low densitypopulations of tigers (Karanth & Chundawat2002), the sample size or the number ofindividuals captured (Mt+1) was small in thisstudy. The statistical test (z = -1.494, P = 0.07)supported the assumption that the tigerpopulation was a closed population during thesampling period. The goodness of fit test for model Mh (as

43

dddd

dd

ddd

dd

ddd d d

dd

d

dd

d

dd

dd

d dd d d

d

ddd

d d

d d dd

d

dd

d

dd

dd

d

ddd

d

001000000000000PAT-111

000001000000000PAT-110

000000000001010PAT-109

000000000000001PAT-108

001000000000000PAT-107

000000000100000PAT-106

101100000001000PAT-105

000000000000010PAT-104

000000000100000PAT-103

000000000011100PAT-102

000000000010000PAT-101

151413121110987654321Individual ID number

001000000000000PAT-111

000001000000000PAT-110

000000000001010PAT-109

000000000000001PAT-108

001000000000000PAT-107

000000000100000PAT-106

101100000001000PAT-105

000000000000010PAT-104

000000000100000PAT-103

000000000011100PAT-102

000000000010000PAT-101

151413121110987654321Individual ID number

Table 9. Capture history of photogrphed tiger in PNP.

Figure 18. Map showing camera trap locations, sampled area (dark green) and non tiger habitat (brown polygon)in PNP.

44

opposed to any alternative model) showed areasonable fit (?2 =15.156; df =14; P = 0.37), asdid a similar test for Model Mb (?2 =24.143; df=19; P = 0.19). Because of sample sizeconstraints, we could not test the fits of modelsMt or Mh against model Mo. The test of thebehavioural response did not provide evidence insupport of model Mb (?2 =1.614; df =1; P = 0.20).Similarly, the test for time-specific variation incapture probabilities during the survey did notprovide support for model Mt (?2 =3.597; df =14;P = 0.997). Further comparisons showed Mh wasa more likely model than Mbh.

The overall model selection test in CAPTUREscored the various competing models as follows:Mo=1.00, Mh=0.91, Mb=0.53, Mbh=0.79,Mt=0.0, Mth=0.46, Mtb=0.49, Mtbh=0.78.Although the null model Mo ranked higher, theestimator based on this model is not robust toviolations of the underlying assumption thatcapture probabilities do not vary amongindividual tigers. Tigers have a complex socialorganization consisting of resident breedersthat maintain home ranges that overlap betweenthe two sexes. Additionally, a proportion of thepopulation consists of non-breeding ‘floaters’that move back and forth across the homeranges of several resident tigers (Sunquist, 1981;Smith, 1993; Karanth & Sunquist, 2000). Thesespatial patterns as well as locations of traps inrelation to tiger movements and home rangeboundaries were likely to cause captureprobabilities to vary among individual tigers.Overall, considering both ecological factors andmodel comparison results, we selected Mh asthe most likely model to represent the underlyingcapture-recapture process that generated thetiger capture histories reported in Table 9. During the survey, 8 individual tigers werecaptured only once, one animal was caught twice,one animal thrice and one animal four times(Table 1). We used the jackknife estimator(Burnham & Overton, 1978; Otis et al., 1978)implemented in CAPTURE, which had givenreasonably acceptable results in earlierphotographic capture studies of tigers (Karanth,1995; Karanth & Nichols, 1998). Average capture probability per sample wasestimated to be 0.0391 by using the Mh jackknifeestimator and the corresponding population sizeestimate ( was 29, (SE= 9.65). Thus, the overallprobability of capturing a tiger was only 38%.Therefore, it was critically important for us tohave used a population estimation method thattook into account the fact that not all animalspresent in the study area would be detected(Nichols, 1992; Williams et al., 2002). The trappolygon encompassed an area of 222.1 km2(Figure 18). The buffer width was estimated tobe 2.94 km, which resulted in 418.14 km2 ofeffective sampled area estimating tiger density.Tiger density was calculated dividing theestimated population size by the effectively

Figure 19. Camera trap photographs of individual tigershowing distinct stripe patterns.

45

sampled area . The density estimated for thesampled area of the Panna Tiger Reserve studyis 6.94 tigers/100 km2.

DiscussionSocial organisation, Population and dispersal The monitoring of seven radio-collared tigersgenerated information on populationcharacteristics, social organization, survival andmortality. This amounts to over 41yearsmonitoring of radio-collared tigers. If theobservation and monitoring of other tigers andcubs is included, it amounts to an astonishing95years of tiger monitoring.

The sex ratio of the 38 tigers that wereidentified positively with sex and age is almostequal to unity (Female19: 20 males). But theratio of the breeding animals favoured thefemales, 3:1 as documented in other studiesconducted in Nepal (Sunqiust, 1981 and Smith,1984). Though the ratio is much smaller thanthat known from more productive habitats whereone male can include up to seven femaleterritories. In 2002, the tiger population in theintensively monitored area consisted of only 31%of breeding individuals, 51% of cubs and subadults and rest of the 17% belonged to nonterritorial, non breeding adults.

Female tigers reach sexual maturity at twoto three years in captivity. There are very fewobservations from the wild. In Chitwan, Sunquist(1981) observed that one radio-collared femalecame into heat at 30 months. In Panna, we hadthe opportunity to radio-collared three youngtigresses at an early age and observed that thefemales gave birth to their first litter on average,at 39.8 months. Considering 100 to 105 daysof gestation period it appears that they wouldcome to heat at about 36 months. The earliestrecord, of 29 months from Panna, is close towhat was observed in Chitwan by Sunquist(1981). This female dispersed earlier then herother two siblings and established herindependent territory first. The other siblingsand cubs produced litters once they settledpermanently in an area. Therefore it appears thatthe time of the first litter is dependent on findinga space for establishing a territory of their own.The average litter size of 2.3 (cubs at the age of2-12 weeks) is very close what is documented by

other field studies in the subcontinent (Schaller,,1967; Sunquist, 1981; Smith, 1984). Anotherlitter is produced after an average period of 21.6months. Observations from other studiesdocument that this period varies considerablyfor different litters from the same animal andbetween different individuals; it can rangeanywhere between 18 to 30 months (Schaller,1967; Sankhala, 1978; Sunquist, 1981; Smith,1984). Surprisingly in Panna, it worked quite clocklike. We observed that almost all the litters (13out of 14 litters) were produced at 22 months’interval, except one which was produced early,within a 20 months gap. This suggests thatfemales come to oestrous when the previouslitter is about 18-19 months old and weaned butnot entirely independent; at this age they arecapable of making their own kills but continue toshare kills with the mother. Observations fromcaptivity suggest a gestation period of 100 to110 days. Based on the observation of twelvebirths and mating average gestation period inPanna is 103 days; this is very similar to what isthe reported period in captivity and from otherfield observations. Mating was seen throughoutthe year; a casual observation may suggestpeaks during winter and monsoon months butour field observations suggest different reasonsfor the oestrous cycle. It appears mainly due tothe almost fixed cycle of oestrous and birth thatwas observed in Panna. In this case it appearsthat the birth of first litter was the determiningfactor and the subsequent mating and birth ofa new litter shifted accordingly by two to threemonths (early).

In this study we were able to monitor andobserve only four dispersals. All female tigerswere radio-collared and established theirterritory at the adjoining vacant spaces and inone case by displacing an old established female.We did not see any evidence of male cubsestablishing a territory in the area. But we sawthem mating with territorial females which werepart of their father’s territory after he had lostits control over the area and it was occupied bya new dominant male.

Estimation of tiger population The results obtained from photographiccapture and recapture method are similar to the

46

To understand the ecology of an animaland its response to the environment, it isimportant to study its populationcharacteristics and ranging pattern, includingits home range size, shape and utilizationpattern. It provides information about spatio-temporal use of its home range over time andseason (Harris et al, 1990). Many authors have tried to define and refinethe concept of home range (Burt, 1943; Mohr,1947; Baker, 1978; Dixon and Chapman, 1980;Anderson, 1982; Worton, 1989). No animalroams at random in an area and each individualhas a home region, where it operates (Seton,1909); the size of home region varies fromspecies to species. In other words, home range/territory can be defined as the area which ananimal uses to carry out its normal activitiesand which fulfils its energy requirement (Burt1943; Jewell, 1966; Brown, 1975; Harris et al.1990).

However, territory is an exclusive use areafully defended by the dominant individual,whereas the home range may be shared either inpart or completely. The animal may use the area

evenly; its use dependant on the ecologicalcharacteristics of the area ? availability anddispersion of its resources, such food, cover andwater. The animal may not patrol 100% of itsterritory regularly; there could be some areas,which it may visit in?frequently (check withoriginal that I have interpreted correctlyuncertain). The importance of such activitycentres (Smith, 1990) is of significantimportance for the management of tigerpopulation as the reproductive success,abundance and viability of a population is largelydependent on these critical habitats/resources.

The size of the home range varies fromspecies to species and it is dependent on bodysize (McNab 1963, Harestad and Bunnell 1979),guild and availability and dispersal of keyresources such as food, water and cover (Seton,1929; Smith, 1990). The home range sizes oftiger and resulting densities are directly relatedto the habitat quality (availability of prey andcover). In prime habitats, which contains highabundance and variety of ungulate prey base, thehome range of a female tiger may be only 20 km2,or even less (Sunquist, 1981; Smith, 1993;

estimation based on the radio-telemetry study.This method which has been used successfullyin many different habitat types (Karanth &Nichols, 1998; 2002) provides a suitablealternative and reliable method for estimatingtiger population over the conventional pug markbased total counts. But it requires careful designand planning to address the issues of spatialsampling and observability (Thompson et al.,1998; Williams et al., 2002). The estimated tiger density of 6.94 animals/100 km2 in Panna is also reasonably high despitehigh anthropogenic pressures and poor ecologicalconditions that exist in a Dry Tropical Forest.These results only indicate the potential of suchtiger habitats in tiger conservation, especiallyin Tropical Dry forests which are the largest tigerhabitat in the country. Tigers in Panna avoid usingthe dry, open forests that form a largeproportion of their home ranges, because of

water-scarcity and lack of shade for thermoregulation although these areas appear tosupport adequate prey densities (R. S.Chundawat, unpubl. data). Therefore, it initiallyappeared that high prey densities might nottranslate into high prey availability for tigers,suggesting that tiger densities may be belowpotential limits set by prey abundance. However,assuming that tigers crop about 10% of standingprey numbers annually (Schaller, 1967; Sunquist,1981; K. U. Karanth, unpubl. data), the estimatedtiger density of 6.94 tigers/100 km2 for Pannais close to the predicted density of 6.18 tigers/100 km2. Therefore, we propose that increasedprey vulnerability and seasonal variations in preyselection behaviour, may enable tigers to achievehigher prey-cropping rates during summer inPanna, despite not fully utilizing their homeranges.

TIGER-II: Home range, movement and activity pattern and habitatuse

47

Karanth and Sunquist, 2000). In the RussianFar East, where the resources are very seasonal,prey availability is low and unevenly distributed,a female may require around 250 to over 400km2 area to fulfil her requirement (Miquelle, etal., 1999).

Chemical Immobilization and Radio-Collaring oftigers:

Like most other cats, the tiger is also anelusive animal, and therefore, it is not alwayspossible to see it. This makes it very difficult tostudy wild free-ranging tigers. To collectecological information in a systematic and un-biased manner one needs to find a tiger at willand this can only be achieved with the help of aradio-collar. A total of 7 tigers were chemicallyimmobilized and captured for the purpose ofradio-collaring.

Chemical Capture The radio collaring of tigers was mostlycarried out in the winter season to avoid theproblem of hyperthermia (rise in bodytemperature due to the effect of drugs).Attempts were made to locate the tigers on kills.Whenever the bait was killed, the area wasscanned from elephant back to locate the tiger.Once the tiger was located, the body weight wasestimated and darts were prepared on the spotby the veterinary doctors. To get the desiredsedation, a combination of Medetomidine (0.05mg/kg body weight) with Ketamine hydrochloride(3.5 mg/kg body weight) was used. Telinjectequipment and pressure syringes were used fordarting. Tigers were darted from elephant backat a distance of about 20-25m. Before dartingtwo to three persons were placed on nearby talltrees to keep a close watch on the tiger’smovement after darting. However, after darting,the tigers did not move much and walked only40-50m distance, or less, before becomingsedated.

After confirming that the animal was fullysedated, it was approached carefully and radio-collared. Information on other parameters suchas body morphometry, its load of ecto-parasitesand its general health condition were recorded.During the operation, a veterinary doctorconstantly monitored the tiger. The entire

operation was carried out in the presence ofthree veterinary experts and finished within 20-25 minutes. The animal was revived with anantidote, Antisedan, a specific antagonist forMedetomidine. Details of the radio-collaredtigers are given in Table

Radio Tracking:For this study, we are using activity radio-

collars, which are manufactured by Telonics Inc.Arizona. The frequency of these collars rangesbetween 150.00 MHz and 151.00 MHz. Theapproximate life of these collars is about 36months and they weigh around 450 gm. Withmovement of the tiger’s neck, the pulse rate ofsignals changes. The radio-collared tigers were tracked withthe help of directional H-antenna, Yagi antennaand a portable receiver. It was observed that theground to ground range of the radio collars wasaround 1.5 to 2.0 km. However, in case of anelevated vantage point the range went up 6.0km. Therefore, hillocks, watch towers and otherelevated places in the field that received thesignals efficiently were identified and marked aspermanent vantage points. Apart from this, theexisting roads were also marked at every 500mto prepare standard reference points. Co-ordinates of such reference points were obtainedusing GPS. Apart from these points, severalother miscellaneous points were generated,whenever required. The radio-collared tigers were tracked fromelephant back, vehicle and at times on foot. Torecord unbiased information, without disturbingthe radio-collared animals, a minimum distanceof 300 to 400m was maintained. Triangulation(Tester and Sniff 1957, Heezen and Tester 1967& Springer 1979) and homing technique (Mech,1983) were used to locate the animals. To getbetter results with minimum error polygon, 4-5bearings were taken from known reference pointsusing the directional antenna and a hand heldcompass.

To obtain the locations (fixes) of thecollared animals, these bearings and values ofstandard reference points were plotted by usinga computer program ‘LOAS’. In case of homing,the locations were directly obtained by usingGeographical Positioning System (GPS). The

48

study area maps were digitized by using GRASS4.1 software and later analysed with the help ofARCHINFO. The locations were then plotted ona 1:50,000 scale Survey of India topo-sheets.Home ranges of each tiger were calculated byusing “Minimum Convex Polygon” (MCP) methodof Mohr (1947). In this method the outer mostlocations are connected to make a polygon andthe area of this polygon gives the size of a homerange. To find out the core areas (areas morefrequently used) of the home range, 55% contourof harmonic mean method (HM) (Dixon andChapman, 1980) was used.

Asymptotes for a number of locations atannual and different seasons were obtained tocheck the adequateness of sample size.

Movement Pattern:To study the activity and movement

pattern of tigers over a period of 24 hours, theywere radio-tracked continuously. This trackingalso gave information on the distribution oflocations in different time zones. Since, radio-tracking tigers for a continuous stretch of 24hrs was tiresome, the cycle was divided into 4phases, each phase comprising of a minimum 6hours. During such tracking phases, the activity(at every 10 minutes) and movement pattern (atevery half an hour) of tigers were recorded.

The information gathered was plotted onthe topographic sheets to study the movementand activity pattern. The seasonal variation inactivity and movement patterns for male andfemale was studied.

ResultsSample size for estimating home rangesThe cumulative home range of radio-collaredtigers is calculated by pooling ten successiveradio-locations. This exercise is done todetermine the minimum number of locationsrequired to estimate a home range size. It isestimated that home ranges of all radio-collaredtigers reached its peak at around 50 locations.In the case of female tigers, the minimum numberof locations required to estimate a home rangesize is between 40 and 50 locations but in thecase of male tigers, it is between 50 and 60locations (Figure XX). At the rate of three tofour locations a week, a minimum of five to sixmonths of monitoring is essential to estimate

a reliable home range size. During this study,every year over 120 locations were obtained foreach of the radio-collared tigers. Therefore, homerange calculations in this study provide a veryreliable estimate.Home range

The radio-collared tigers were monitoredon a regular schedule as described in themethods. Home ranges are estimated using thesoftware programmes “BIOTAS” and “LOAS”(XXXXX). Although information on the activity andmovement patterns for all the radio-collaredanimals is recorded, the data is not included inthis report. During the course of the study, seventigers were radio-collared but only six of themwere monitored for more than six months andinformation on these animals are discussed indetail in this report. These animals include fourfemales and two males; details of the radio-tracking is provided in table 10a. The first animalthat was radio collared was a sub-adult female,which died within five months of tracking due tounknown causes. Male tiger-91 was the secondtiger radio-collared in the study followed byTigress-118. Her three daughters from the samelitter were subsequently collared and weremonitored for several years. In this report datafrom 1997 to 2002 is used for describing homerange and movement patterns.

Annual home ranges The annual home ranges of males were fourand half to five times larger than female ranges.The average annual range calculated for maletigers is 184 km2. The area of most activity(80%), which can be defined as the core area, onaverage remained limited to less than 20% ofthe average annual home range. This remainedmore or less consistent throughout the studyperiod in both the males, despite considerablevariation seen in the annual home range sizes indifferent years. Average annual home range offemales is estimated to be 42.16 km2, whichranged between 24 km2 and 83.6 km2. The corearea, which represents 80 % of their activities,remained consistently on average within 20% ofthe annual home range. The home ranges duringthe full monitoring period is considerably largerthen the average annual home range (Table 10).The average home range size for the study period

49

0

50

100

150

200

250

300

10 30 50 70 90 110

130

150

170

190

210

230

No. Location

Sq

. K

ilo

met

res

Fm-111 Fm-118 Fm-120 M-91

24 months 220-23096 –100 months12-01-2002-M-125

22 months56 months21-12-2000Fm-113

53 months24072-80 months 11-04-1996 andOct. 1998

M-91

42 months11534 months 07-01-1999 and27-01-2001

Fm-120

52 months10824 months26-03-1998 and22-12-2000

Fm-111

30 months12050-55 months27-01-1997 to Nov. 199726-01-2001 and10-1-2002

Fm-118/123

Six monthsSub-adult09-04-1996Fm01

Tracking Period(In Months)

Weight (Kg.)

Age at the time of collaring

Capture DateTiger No.

24 months 220-23096 –100 months12-01-2002-M-125

22 months56 months21-12-2000Fm-113

53 months24072-80 months 11-04-1996 andOct. 1998

M-91

42 months11534 months 07-01-1999 and27-01-2001

Fm-120

52 months10824 months26-03-1998 and22-12-2000

Fm-111

30 months12050-55 months27-01-1997 to Nov. 199726-01-2001 and10-1-2002

Fm-118/123

Six monthsSub-adult09-04-1996Fm01

Tracking Period(In Months)

Weight (Kg.)

Age at the time of collaring

Capture DateTiger No.

Table 10a. Date of capture, age at the time of callaring and duration of radio-tracking of radio-collared tigers

Figure 20a. Estimation of minimum nuber of radio location required to estimate a reliable homerange size for different radio-collared tigers.

5024.313.153.8Annual average

15.03.624.02002.0

27.022.683.62001.0

19.616.483.6Total 2001-02

Femlale-113


16.76.739.92002.0

28.312.845.32001.0

26.79.836.72000.0

32.312.538.81999.0

31.216.954.2Total 1999-2002

Female-120


17.84.726.22002.0

13.73.626.32001.0

27.013.750.52000.0

13.34.634.91999.0

20.212.360.91998.0

31.416.653.0Total 1999-2002

Female-111


28.49.232.32001-02

21.88.237.52000-01

34.213.940.6Total 2000-02

32.813.541.01997.0

Female-118

19.436.6188.6Males- Annualaverage

13.628.6210.0Male-125

0.091.02000.0

20.327.9137.11999.0

23.843.8184.11998.0

25.164.6257.01997.0

21.754.8252.21996.0

23.568.3290.0Total 1996-2001

Male-91

% use

Harmonic Mean

Minimum Convex Polygon

Radio-collared Tigers


15.03.624.02002.0

27.022.683.62001.0

19.616.483.6Total 2001-02

Femlale-113


16.76.739.92002.0

28.312.845.32001.0

26.79.836.72000.0

32.312.538.81999.0

31.216.954.2Total 1999-2002

Female-120


17.84.726.22002.0

13.73.626.32001.0

27.013.750.52000.0

13.34.634.91999.0

20.212.360.91998.0

31.416.653.0Total 1999-2002

Female-111


28.49.232.32001-02

21.88.237.52000-01

34.213.940.6Total 2000-02

32.813.541.01997.0

Female-118

19.436.6188.6Males- Annualaverage

13.628.6210.0Male-125

0.091.02000.0

20.327.9137.11999.0

23.843.8184.11998.0

25.164.6257.01997.0

21.754.8252.21996.0

23.568.3290.0Total 1996-2001

Male-91

% use

Harmonic Mean

Minimum Convex Polygon

Radio-collared Tigers

Table 10. Over all and annual home ranges of radio-collared tigers in PNP.

51

66.627.347.135.562.828.442.7Femaleaverages

77.130.939.215.778.531.540.2average

87.534.920.38.162.825.139.92002

65.529.642.419.282.337.345.32001

65.624.154.720.185.831.536.72000

89.734.839.315.383.232.338.81999

F-120

72.927.070.426.562.522.736.9average

74.023.948.915.879.525.732.32002

59.022.183.631.353.720.137.52001

85.635.178.832.354.422.341.01997

F-118

69.534.237.420.348.834.353.8average

80.319.336.78.821.95.324.02002

58.849.138.131.875.863.383.62001

F-113

47.017.241.315.461.525.039.8average

88.923.349.212.971.018.626.22002

38.010.036.19.524.46.426.32001

61.030.834.817.681.341.150.52000

26.69.358.720.579.027.634.91999

20.512.527.416.751.831.560.91998

F-111

%Winter%Summer%MonsoonYearlyFemales

55.098.969.7124.645.174.7184.0average

210.0M-125-2002

82.9212.146.3118.539.8102.0256.01996.0

29.273.666.3167.218.245.9252.01997.0

37.468.996.2177.140.274.0184.01998.0

56.076.771.798.277.5106.2137.01999.0

69.763.468.161.949.845.391.02000.0

M-91

% of annual

Winter% of annual

Summer% of annual

MonsoonYearlyTiger IDs

66.627.347.135.562.828.442.7Femaleaverages

77.130.939.215.778.531.540.2average

87.534.920.38.162.825.139.92002

65.529.642.419.282.337.345.32001

65.624.154.720.185.831.536.72000

89.734.839.315.383.232.338.81999

F-120

72.927.070.426.562.522.736.9average

74.023.948.915.879.525.732.32002

59.022.183.631.353.720.137.52001

85.635.178.832.354.422.341.01997

F-118

69.534.237.420.348.834.353.8average

80.319.336.78.821.95.324.02002

58.849.138.131.875.863.383.62001

F-113

47.017.241.315.461.525.039.8average

88.923.349.212.971.018.626.22002

38.010.036.19.524.46.426.32001

61.030.834.817.681.341.150.52000

26.69.358.720.579.027.634.91999

20.512.527.416.751.831.560.91998

F-111

%Winter%Summer%MonsoonYearlyFemales

55.098.969.7124.645.174.7184.0average

210.0M-125-2002

82.9212.146.3118.539.8102.0256.01996.0

29.273.666.3167.218.245.9252.01997.0

37.468.996.2177.140.274.0184.01998.0

56.076.771.798.277.5106.2137.01999.0

69.763.468.161.949.845.391.02000.0

M-91

% of annual

Winter% of annual

Summer% of annual

MonsoonYearlyTiger IDs

Table 11. Seasonal home ranges of the radio-collared tigers in PNP.

52

0.465 km4.37 ± 0.26 kAnnual

2.296 km4.53 ± 0.48 Winter

3.664 km4.74 ± 0.74Summer

2.203 km3.85 ± 0.49Monsoon

M-91

0.238 km2.11 ± 0.13Annual

0.257 km2.04 ± 0.14 Winter

0.871 km1.92 ± 0.50 Summer

2.232 km2.38 ± 1.00 Monsoon

F-120

0.309 km1.73 ± 0.17Annual

0.887 km1.58 ± 0.44Winter

0.947 km1.53 ± 0.47Summer

1.462 km2.09 ± 0.65Monsoon

F-111

0.045 km1.7 ± 0.026Annual

1.169 km1.71 ± 0.52Winter

0.905 km1.65 ± 0.286Summer

1.089 km1.74 ± 0.36Monsoon

F-123

Std. Dev.Mean Distance km (SE)

Animal

0.465 km4.37 ± 0.26 kAnnual

2.296 km4.53 ± 0.48 Winter

3.664 km4.74 ± 0.74Summer

2.203 km3.85 ± 0.49Monsoon

M-91

0.238 km2.11 ± 0.13Annual

0.257 km2.04 ± 0.14 Winter

0.871 km1.92 ± 0.50 Summer

2.232 km2.38 ± 1.00 Monsoon

F-120

0.309 km1.73 ± 0.17Annual

0.887 km1.58 ± 0.44Winter

0.947 km1.53 ± 0.47Summer

1.462 km2.09 ± 0.65Monsoon

F-111

0.045 km1.7 ± 0.026Annual

1.169 km1.71 ± 0.52Winter

0.905 km1.65 ± 0.286Summer

1.089 km1.74 ± 0.36Monsoon

F-123

Std. Dev.Mean Distance km (SE)

Animal

Female-118 occupies an undisturbed highprey density habitat in the core area of the Park.The estimated home range for 2000 to 2002(MCP 100%) is 40.6 km2 with the core area 13.9km2 - where she spends over 80% of the time;this represents only 34% of the range. Her rangeoverlapped with that of her daughters, female-111, female-120 and female-113 (Figure 20).However, we did not see any association withher daughters after they became independentand established their territories. Soon overlapwith the daughters’ territories reducedconsiderably. Female-118 had a larger home rangeprior to her first litter establishing their

for female is 57.85 km2 and for males it is210 km2.

Seasonal home rangesOn average seasonal home ranges are

small and represented 40-65 % of the annualhome ranges. No clear seasonal trend in homerange size is observed in radio collared tigersof the Tiger Reserve. Seasonal ranges, however,changed considerably in size over the years.For detail see table 11.

Monitoring Radio-collared TigersFemale-118

Tigress ‘118’ was radio-collared inJanuary 1997 (table-10a) and was monitoredfor eleven months. Her collared stoppedfunctioning in November 1997. The lastlocation obtained from that collar was on29th November 1997. The research team wasable to monitor her without her collar, but in alimited way. Several attempts were made toreplace her old collar but her non-functionalcollar was replaced only in February 2000. InDecember 2000, while dragging a sambar kill,this collar broke off. She was radio-collaredfor the third time within a month, in January2002. Her last location was recorded duringMarch 2005 after 38 months of monitoringby the radio-collar. The female-118 was raisinga litter of three 7-8 month-old female cubs,at the time she was collared for the first time.Unfortunately the radio-collar malfunctionedafter eleven months. The research teamcontinued to monitor her movements andother activities, though intermittently, till shewas radio collared once again. Later the researchteam was able to radio-tag all the three femalecubs of her first litter after they becameindependent and had established their ownterritories. In the in-between period of the firstand second collaring, Female-118 had a secondlitter of two male cubs, which, like her first litter,survived to adulthood and dispersed. At the timeof the second radio-collaring she was raising herthird litter of three cubs comprising of two malesand one female. Her third litter also survived toadulthood and dispersed. She gave birth to afourth litter comprised of two males and onefemale.

Table 11. Average distance travelled by radio-collared tigersin 24 hours.

53

territories. Subsequently part of her range wastaken over by her daughters and as a result herrange reduced by approximately 20%. Her annualhome range size is 34.9 km2 and 80% of heractivity is limited to 25% of the annual homerange. Home ranges changed with the seasonand also over the years (see Table-11 for details).Her monsoon ranges (22.7 km2) are smaller thanher summer (26.5 km2) and winter (27 km2)ones. Seasonal home ranges of tigress-118 onlyaccount for 60 to 70 % of the annual homeranges (Figure 30).

Female-111Female-111 is one of the three siblings of

the first litter of female-118. She was first radio-collared in April 1998, when she was in theprocess of dispersing. Initially she used part ofthe mother’s territory frequently and stayedthere for several months before establishing herown territory on the upper Talgaon plateau. After6-7 months of several forays into thesurrounding areas, she established her ownterritory in an adjoining area immediatelysoutheast of her mother with some overlap. Herterritory extended beyond the Park boundary bythree to four kilometres into the neighbouringterritorial forests of South Panna ForestDivision. She had her first litter of two cubs -one male and one female - at the age ofapproximately 36 months, in May 1999. She hadher second litter in March 2001, twenty-twomonths later. The second litter comprised ofthree cubs but the research team was not ableto identify the sex of these cubs. Female-111occupied an area close to the periphery of theTiger Reserve and her territory included areasclose to several villages situated inside and atthe periphery of the Reserve. The research teamlost contact with her in September 2002. Herlast locations were on a cattle kill and shedisappeared mysteriously after feeding for acouple days on the kill. The research team madeattempts to confirm her presence by placingcamera traps in the area and monitoring herfrequently used areas but failed to find anyevidence even after three months of intensivemonitoring. Later, after six months, this emptyterritory was occupied by another tigress, whowas seen mating with the male tiger-125 and

photographed. This confirmed that tigress-111had most likely died due to unknown causes, butfrom circumstantial evidence and because shedisappeared from a cattle kill, it is reasonableto assume that it is possible that she was victimof human caused mortality.

She was radio-tracked for fifty fourmonths (4.5 years). Female-111 patrolled a largearea of 53 km2 (100% MCP)and most of heractivity (80% HM) was restricted within 31% ofher range on the upper Talgaon plateau (Table10). The area used by her has low prey abundancebut very high cattle presence and most of herkills were of cattle. Moreover, during the hotsummer months, when there is no water availablewithin her territory on the upper plateau insidethe Tiger Reserve, she either has to come to thebase of the escarpments on the Hinauta plateauin search of cool resting-places and water or hasto visit a water hole more then 3 km outside theReserve boundary (Figure 22). In this area sheregularly killed livestock. This brought her intodirect conflict with local human populations,whenever she killed livestock in this unprotectedforest habitat. Her average annual home rangesize calculated is 39.8 km2. This highlights thefact though she used part her home range overthe years but her annual range and centre ofactivity changed over the years (Table 11). Thatis reflected in her small centre of activity (6.6km2) representing only 16.7% of the annual rangeon average. In contrast to tigress-118, female-111’s monsoon home range was larger (25 km2)than her summer (15.4 km2) and winter (17.2km2) ones. Seasonal ranges represent 40 to 60% of the annual ranges (Figure 29).

Female-120Female-120 was radio-collared in January

1999, when she was approximately 32 monthsold. She had established her territory in the areawest of her mother’s territory, which sheoccupied by displacing an old territorial femalefrom the area (Figure 25). She was able todisplace the territorial female with the help ofher third sibling (female-113), who lived and movedin coalition with her for eight to nine monthsbefore they separated and established theirrespective territories. At the time of her collaringshe was accompanied by her sibling and remained

54

in association with her for another couple ofmonths after radio collaring. She gave birth toher first litter at the age of around 42 monthsin December 1999. This litter comprised of twocubs, one male and one female. These cubs grewto adulthood and dispersed successfully. Shegave birth to her third litter in 2000. During thistime the dominant male-91 was replaced byanother male tiger we called ‘Hairy foot’. Hermovements were different from her previous yearand she never brought her litter to her core areafor any duration; this area was frequentedregularly by ‘Hairy foot’. She was constantly onthe move and her behaviour indicated stress. Welast saw her litter in December 2001. She gavebirth to a third litter in March 2002, whichconfirmed our doubts that ‘Hairy foot’ had killedher second litter, fathered by male-91. What wasinteresting was that female-120 apparentlycame into oestrus and mated successfully whenher second litter were still alive. This is the firstrecord from the wild of a female coming intooestrus while raising a litter and giving birth toa new litter within 100 days of losing the litterto infanticide. ‘Hairy foot’ was seen on severaloccasions visiting the litter site and spentseveral hours with the female within a few metresof weeks-old cubs at regular intervals.Unfortunately, ‘Hairy foot’ did not survive longand was found dead in mysteriouscircumstances in a well inside the Reserve. Withina few months, this exposed the female-120 yetagain to another male tiger who had not fatheredher cubs. She once again left her core area andmoved into an area away from the new male’sactivity. This area, at the periphery of park,suffers from intense human pressure. One tigerwas poached in this area in the early nineties,well before the project began. Female-120succumbed to the same fate as the earlier tigerand was found dead in a snare. At the time ofher death, her two cubs were almost nine monthsold. Like her mother, she also occupied a prey-rich, undisturbed forest habitat on the middleplateau. But her range extended up to the Parkboundary and included areas adjoining theterritorial forest and villages at the southernperiphery of the Park. She was not as successfulas her mother and other siblings as she was

exposed to a frequent change-over of males inthe area, which coincided with litters fatheredby previous males. She was radio tracked for 35 months; duringthis period she ranged over an area of 54.2 km2(MCP 100%). Most of her activity was restrictedto small areas of 16.9 km2, which representabout 31% of the total range (Table 10). Heraverage home range size is calculated as 40.2km2 and its centre of activity (80% HM) islimited to 10 km2 only. Seasonal ranges are smallbut her monsoon and winter ranges are relativelylarge and represent over 70% of the annualranges. Summer range is smallest approximately40% of the average annual range (Figure 27).

Female-113The third daughter of female-118, was

radio-collared in December 2000. Like her othertwo siblings (F-111 and F-120), she also hadestablished a territory in an area adjoining themother’s territory and partially occupying female118’s range. Her territory was to the north ofher mother’s, female-118’s range (Figure 23). Atthe time of her collaring in December 2000, shewas also raising a litter of two cubs, one maleand a female, 15-16 months old. She occupiedan area on either side of the River Ken but mostof her range included areas south of the riverand in the tourism zone. Her range also extendedup to the periphery of the Reserve north of theKen river. Her collar was removed in 2002because of disturbance created by heavypressure from tiger shows that occurred almostevery day for several hours during the peaktourist season. She later disappeared in 2004and was last seen in December 2003. She hadraised two litters to adulthood, which dispersedsuccessfully.

She covered a large territory of 83 km2(MCP 100%) but spent most of her time in amuch smaller core area of 16.4 km2 (Harmonicmean 80%) which is of similar in size to thatestimated for her other siblings? but not female-118; Table 11. Her range reduced to 24 km2 duringthe second year with a core area of only 3.6 km2.She occupied a very narrow belt of tiger habitatalong the river Ken. Her seasonal range sizechanged considerably across the seasons andover the years too. On average, the winter ranges

55

showed the largest area patrolled by the tigress,representing almost 69% of the annual range;this was followed by that of the monsoon rangeof 48.8% (Figure 28).

Male-91The male-91 was radio-collared in April

1996; its first collar was replaced in October1998. His second collar continued to give signalfor the next two years and finally stoppedfunctioning in October 2000. At its peak, male-91’s range covered an area of 290 km2, which ismore then 50% of the entire Park area (table 11).At the early stages of monitoring, in 1996 and1997, this enormous range included only twofemale territories female-118 and the other,Bargad,i female, who was later ousted by female-120. Later, between 1998 and 2000, his territoryincluded 118’s daughters, female-120 and female-111 but he lost part of his range to male-125;this part included the territory of the thirdsibling, female-113. This occurred after a seriousfight that took place in September 1998. Withfemale-118, 111 and 120 he fathered six littersand two from the Bargadi female. These eightlitters amount to seventeen cubs, of whichthirteen survived to adulthood in the five and ahalf years when he remained as dominant malein the area. Over the years he has lost aconsiderable part of his territory to other males,125 and ‘Hairy foot’, after several battles. Histerritorial behaviour has been discussed in detailin earlier reports and publications (Chundawatet al, 1999). After these territorial fights hisrange was reduced to a mere 91 km2 and onlypartially covered the territories of the threefemales he once dominated (Figure 21). His annual range gradually declined from over250 km2 to 91 km2 at the end of his tenure adominant male of the area.

Male-125 The tiger, male-125 was radio-collared inJanuary 2002; therefore only information fromthe first year is discussed in this report. Male-125 first appeared in mid 1998 in the Madla areaand later he was seen fighting with Male-91. Thisbattle resulted in some very serious injuries toM-91. As a result, male-125 took over a largepart of male-91’s area. Over the next few years,

male 125 extended his range across all threeplateaux, covering an area from Billiya Seha toareas adjoining Rampura. His territory includedthree females’ territories and one of them,female-113, was radio-collared. The other twowere a female on the upper plateau, in the areabetween Jardhoba and Rampura village known asthe ‘Amdar female’ and one that occupied arange around the Gata area and known to us asthe ‘Gata female’.

After the death of ‘Hairy foot’ male-125partially moved into ‘Hairy foot’s territory, butafter mating with female-113 for the third timein 2003 he left the area and remained exclusivelyon the upper Talgaon and Hinauta plateaux. Ayounger male, ‘Broken tooth’ was seen in the areaand soon after male-125 left the area, this youngmale also mated with female-113, which unusuallycontinued for several weeks. In 2003 and 2004,their activities became somewhat complicated:male-125 returned to patrol the area in 2004and ‘Broken tooth’ left it and was seen laterpatrolling the majority of the territory once usedby ‘Hairy foot’. Female-113 disappeared and hermother, female-118, extended her range and re-occupied part of this vacant space; this wasafter six years and beyond her original territory.Her doin g this further confirmed the loss offemale-113 from the area. In this area male-125was seen frequently in association with female-118. This was the first time since we knew theseindividuals that they were seen together.

Male-125 occupied a fairly large territory of 210km2 in the year 2002. Most of his activityremained within 20% of his annual range, whichis similar to that of male-91 (Figure 26).

Movement Pattern The average daily distance travelled by femaletigers is 1.84 km, The distance is calculated asthe distance between two locations onsuccessive days and these were taken 24 hoursapart. This information only provides anapproximate index of movement by the radio-collared animals but does not provideinformation on the actual distances covered bythe animals. Actual distances travelled by theanimals in a 24-hour period are much larger thanthe distances calculated in a straight line

56

Figu

re 2

0. H

ome

rang

es o

f ra

dio-

colla

red

tig

ers

in P

anna

Nat

iona

l Par

k.

57

Figu

re 2

1.nn

ual h

Hom

e ra

nges

of

mal

e-09

1 r

adio

-col

lare

d t

iger

in

Pan

na N

atio

nal P

ark.

58

Figu

re 2

2. A

nnua

l ho

me

rang

es o

f ra

dio-

colla

red

tig

er f

emal

e-11

1 in

Pan

na N

atio

nal P

ark.

59

Figu

re 2

3. A

nnua

l ho

me

rang

es o

f ra

dio-

colla

red

tig

er f

emal

e-11

3 in

Pan

na N

atio

nal P

ark.

60

Figu

re 2

4. A

nnua

l ho

me

rang

es o

f ra

dio-

colla

red

tig

er f

emal

e-11

8/13

in

Pan

na N

atio

nal P

ark.

61

Figu

re 2

5. A

nnua

l ho

me

rang

es o

f ra

dio-

colla

red

tig

er f

emal

e-12

0 in

Pan

na N

atio

nal P

ark.

62

Annaul range

Winter range

Monson range

Summer range

Male-91Seasonal Home Ranges

1999Annaul range

Winter range

Monson range

Summer range


1999Annaul range

Winter range

Monson range

Summer range


1999Annaul range

Winter range

Monson range

Summer range


1999Annaul range

Winter range

Monson range

Summer range

Annaul range

Winter range

Monson range

Summer range


1999

Figure 25. Seasonal home ranges of radio-collared tiger male-0091 in Panna National Park.

63

Annual range

Winter range

Monson range

Summer range


2002Annual range

Winter range

Monson range

Summer range


2002Annual range

Winter range

Monson range

Summer range


2002Annual range

Winter range

Monson range

Summer range

Annual range

Winter range

Monson range

Summer range


2002

Figure 26. Seasonal home ranges of radio-collared tiger male-125 in Panna National Park.

64

Annaul range

Winter range

Monson range

Summer range

Female-120Seasonal Home Ranges

2000Annaul range

Winter range

Monson range

Summer range


2000Annaul range

Winter range

Monson range

Summer range


2000Annaul range

Winter range

Monson range

Summer range

Annaul range

Winter range

Monson range

Summer range


2000

Figure 27. Seasonal home ranges of radio-collared tiger female-120 in Panna National Park.

65

Annaul range

Winter range

Monson range

Summer range


2002Annaul range

Winter range

Monson range

Summer range


2002Annaul range

Winter range

Monson range

Summer range


2002Annaul range

Winter range

Monson range

Summer range

Annaul range

Winter range

Monson range

Summer range


2002

Figure 28. Seasonal home ranges of radio-collared tiger female-113 in Panna National Park.

66

Annaul range

Winter range

Monson range

Summer range


2001Annaul range

Winter range

Monson range

Summer range


2001Annaul range

Winter range

Monson range

Summer range


2001Annaul range

Winter range

Monson range

Summer range

Annaul range

Winter range

Monson range

Summer range


2001

Figure 29. Seasonal home ranges of radio-collared tigerfe male-111 in Panna National Park.

67

Annaul range

Winter range

Monson range

Summer range

Female-118/123Seasonal Home Ranges

2001Annaul range

Winter range

Monson range

Summer range


2001Annaul range

Winter range

Monson range

Summer range


2001Annaul range

Winter range

Monson range

Summer range

Annaul range

Winter range

Monson range

Summer range


2001

Figure 30. Seasonal home ranges of radio-collared tiger female-118/13 in Panna NationalPark.

68

between two locations. Details for individualanimals are given in Table 12. Male tigers covereddistances twice as large as the female tigers.The average distances travelled by male tigersis estimated at 4.37 km. No clear seasonalpattern is observed in distances travelled bytiger in Panna Tiger Reserve, which is very similarto the seasonal home range size observed in theReserve.

Activity PatternTigers in Panna Tiger Reserve show distinct

seasonal activity patterns. On average they arecrepuscular, showing two activities bursts(defined as more than fifty percentactive records). This pattern isclearer during the monsoonmonths showing two distinctactivity peaks – one in the earlymorning hours, between 0600 to0800 hours, and second in theevening between 1800 to 2100hours. During the middle of the dayactivity reaches its minimum.However, in winter only one distinctpeak of activity is observed: in earlymorning hours 0700 to 1000 andhovering around fifty percent in theevening from 1600 to 2100 hours.Activity declines to its minimumduring the middle of the night andremains below 30% throughout thenight or until sunrise. Activity picksup with sunrise and remains over30% for most of the day hours. Thesummer activity pattern is insharp contrast to the winterpattern: only one distinct activitypeak is seen where it reaches above50% after sunset around 1900.Activity hovers around 40% duringthe most of the 24 hour period butreaches its minimum and below20% during midday between 1100and 1700 hours (Figure 31).

Habitat UseHabitat use by tiger

All tiger locations are pooledinto male and female groups and

analysed to determine habitat use by tiger. Adetailed map of different habitat types isprepared from satellite data (Figure 1). From thisdetailed habitat map, availability of each habitatis calculated for tigers. Six different habitattypes are identified and mapped:1. Disturbed (including human habitation andhighly degraded habitat).2. Open/Grassland (including grassland and veryopen savanna type forest habitat).3. Thorn forest4. Miscellaneous5. Mixed forest6. Mixed dense Forest

Monsoon

010

2030405060

7080

1 3 5 7 9 11 13 15 17 19 21 23

Hours

Per

cen

t A

ctiv

ity

Winter

0

1020304050607080

1 3 5 7 9

11

13

15

17

19

21

23

Hours

Per

cen

t Ativ

ity

Summer

0

20

40

60

80

100

1 3 5 7 9 11 13 15 17 19 21 23

Hours

Per

cen

t A

ctiv

ity

Monsoon

010

2030405060

7080

1 3 5 7 9 11 13 15 17 19 21 23

Hours

Per

cen

t A

ctiv

ity

Winter

0

1020304050607080

1 3 5 7 9

11

13

15

17

19

21

23

Hours

Per

cen

t Ativ

ity

Summer

0

20

40

60

80

100

1 3 5 7 9 11 13 15 17 19 21 23

Hours

Per

cen

t A

ctiv

ity

Winter

0

1020304050607080

1 3 5 7 9

11

13

15

17

19

21

23

Hours

Per

cen

t Ativ

ity

Summer

0

20

40

60

80

100

1 3 5 7 9 11 13 15 17 19 21 23

Hours

Per

cen

t A

ctiv

ityFigure 31. Seasonal activity chart of the radio-collared tigers inPNP.

69

X2=608.080810546875

Z= 2.7299993038177

0.015 * -0.0041980.80.04Water bodies

0.345 * +0.288635392.30.196Dense Forest

0.181 * +0.136318202.10.101Mixed Forest

0.329 * +0.273604528.30.264Miscellaneous

0.049 * -0.02675191.20.095Thorn Forest

0.155 * -0.114270445.70.222Open/Grassland

0.028 * -0.01139153.70.077Disturbed

0.031 * +0.0134510.80.005Outside

UpperCL

LowerCL

Observed Usage

Expected usage

Relative area

Habitat categories

a. Female

X2=608.080810546875

Z= 2.7299993038177

0.015 * -0.0041980.80.04Water bodies

0.345 * +0.288635392.30.196Dense Forest

0.181 * +0.136318202.10.101Mixed Forest

0.329 * +0.273604528.30.264Miscellaneous

0.049 * -0.02675191.20.095Thorn Forest

0.155 * -0.114270445.70.222Open/Grassland

0.028 * -0.01139153.70.077Disturbed

0.031 * +0.0134510.80.005Outside

UpperCL

LowerCL

Observed Usage

Expected usage

Relative area

Habitat categories

a. Female

Table 13 Habitat use by all female (a) and male (b) radio collared tigers in PannaTiger Reserve. (* = significant usage; + = preferred; - = avoided)

x=643.3832397460938

Z= 2.72999930381774

0.015 * -0.0041980.80.04Water bodies

0.345 * +0.288635392.30.196Dense Forest

0.181 * +0.136318202.10.101Mixed Forest

0.329 * +0.273604528.30.264Miscellaneous

0.049 * -0.02675191.20.095Thorn Forest

0.155 * -0.114270445.70.222Open/Grassland

0.028 * -0.01139153.70.077Disturbed

0.031 * +0.0134510.80.005Outside

Upper CL

LowerCL

Observed Usage

Expected usage

Relative Area

Habitat categories

b. Male

x=643.3832397460938

Z= 2.72999930381774

0.015 * -0.0041980.80.04Water bodies

0.345 * +0.288635392.30.196Dense Forest

0.181 * +0.136318202.10.101Mixed Forest

0.329 * +0.273604528.30.264Miscellaneous

0.049 * -0.02675191.20.095Thorn Forest

0.155 * -0.114270445.70.222Open/Grassland

0.028 * -0.01139153.70.077Disturbed

0.031 * +0.0134510.80.005Outside

Upper CL

LowerCL

Observed Usage

Expected usage

Relative Area

Habitat categories

b. Male

70

Z= 2.4999995231628

x=456.992156982421

0.282 * +0.2335172770.138High

0.283 * +0.2345183270.163Medium

0.468 * -0.4128821304.00%0.651Low

0.0550.03288960.048Very low

UpperCL

LowerCL

Observed Usage

Expected Usage

Relative Area

Chital abundance

a. Female tiger

Z= 2.4999995231628

x=456.992156982421

0.282 * +0.2335172770.138High

0.283 * +0.2345183270.163Medium

0.468 * -0.4128821304.00%0.651Low

0.0550.03288960.048Very low

UpperCL

LowerCL

Observed Usage

Expected Usage

Relative Area

Chital abundance

a. Female tiger

Table 14 Use of space in relation to chital abundance by all female (a) andmale (b) radio collared tigers in Panna Tiger Reserve. (* = significant usage; += preferred; - = avoided).

x=292.447875976562

Z= 2.4999995231628

0.280 * +0.217291161.80.138High

0.290 * +0.226302191.00.163Medium

0.450 * -0.378485762.10.651Low

0.099 * +0.069356.20.048Very low

UpperCL

LowerCL

ObservedUsage

Expected Usage

RelativeArea

Chital abundance

B. Male tiger

x=292.447875976562

Z= 2.4999995231628

0.280 * +0.217291161.80.138High

0.290 * +0.226302191.00.163Medium

0.450 * -0.378485762.10.651Low

0.099 * +0.069356.20.048Very low

UpperCL

LowerCL

ObservedUsage

Expected Usage

RelativeArea

Chital abundance

B. Male tiger

71

x= 456.9921569824

Z= 2.499999523162

0.461 * +0.405868448.3180.224High

0.042 * -0.0236591.2280.045Medium

0.1550.117272236.590.118Low

0.426 * -0.3728001228.66%0.613Very low

UpperCL

LowerCL

Observed Usage

Expected usage

Relative Area

Sambar abundance

a. Female tiger

x= 456.9921569824

Z= 2.499999523162

0.461 * +0.405868448.3180.224High

0.042 * -0.0236591.2280.045Medium

0.1550.117272236.590.118Low

0.426 * -0.3728001228.66%0.613Very low

UpperCL

LowerCL

Observed Usage

Expected usage

Relative Area

Sambar abundance

a. Female tiger

Table 15 Use of space in relation to sambar abundance by all female (a) and male(b) radio collared tigers in Panna Tiger Reserve. (* = significant usage; + = preferred;- = avoided).

x=335.470642089843

Z= 2.49999952316284

0.475 * +0.403514261.8360.224High

0.0760.0426953.280.045Medium

0.1220.078117138.1780.118Low

0.438 * -0.366471717.5890.613Very low

UpperCL

LowerCL

Observed Usage

Expected Usage

Relative Area

Sambar abundance

b. Male tiger

x=335.470642089843

Z= 2.49999952316284

0.475 * +0.403514261.8360.224High

0.0760.0426953.280.045Medium

0.1220.078117138.1780.118Low

0.438 * -0.366471717.5890.613Very low

UpperCL

LowerCL

Observed Usage

Expected Usage

Relative Area

Sambar abundance

b. Male tiger

72

x=349.1095275

Z=2.499999523162

0.418 * -0.363783984.250.491High

0.254 * +0.207463336.80.168Medium

0.338 * +0.286625376.90.188Low

0.081 * -0.053134306.90.153Very low

UpperCL

LowerCL

Observed Usage

Expected Usage

Relative Area

Nilgai abundance

a. Female tiger

x=349.1095275

Z=2.499999523162

0.418 * -0.363783984.250.491High

0.254 * +0.207463336.80.168Medium

0.338 * +0.286625376.90.188Low

0.081 * -0.053134306.90.153Very low

UpperCL

LowerCL

Observed Usage

Expected Usage

Relative Area

Nilgai abundance

a. Female tiger

Table 16 Use of space in relation to nilgai abundance by all female (a) and male (b)radio collared tigers in Panna Tiger Reserve. (* = significant usage; + = preferred; -= avoided).

x=211.211044311523

Z= 2.4999995231628

0.391 * -0.321417574.80.491High

0.272 * +0.21282196.70.168Medium

0.348 * +0.28368220.10.188Low

0.110 * -0.068104179.20.153Very low

UpperCL

LowerCL

Observed Usage

Expected Usage

RelativeArea

Nilgai abundance

b. Male tiger

x=211.211044311523

Z= 2.4999995231628

0.391 * -0.321417574.80.491High

0.272 * +0.21282196.70.168Medium

0.348 * +0.28368220.10.188Low

0.110 * -0.068104179.20.153Very low

UpperCL

LowerCL

Observed Usage

Expected Usage

RelativeArea

Nilgai abundance

b. Male tiger

73

7. Water bodies

Most of the Tiger Reserve areas(approximately 32%) fall under open foresthabitat, i.e. open/grassland and thorn forest.Approximately 29 % of the area is covered byrelatively dense forest and 26 % is mediumdensity forest. On average, tigers used thedensely forested areas more then other habitatsand of these, the mixed and mixed dense Forestalong the base of the escarpment was used most(female, 47% and male, 52%, Figure 32). Whenthe same data is looked at in more detail withinformation segregated for the summer andwinter seasons, it is noticed that open habitatsare used relatively more (8-15%) in winter.However, overall, distribution of tigers during thewinter months is more widely spread across thehabitats with higher use of dense forest (< 25%).During the summer months it is the mixed foresthabitats along the escarpment that are usedmost. These forest habitats along theescarpments provide special micro-habitats withdense cover, water and caves for shelter, wherethe tiger finds cool places to escape from theintense heat of the hot summer days.

Space use by tiger in relation to its preyavailability The study aims to find out whether thedistribution of tiger is related to its prey and ifit is then to which prey species it is more closelyrelated. A distribution map of densities for majorprey species (these include only sambar, chitaland nilgai) was developed with the help of fielddata and GIS. These models were then taken intothe field for ground truthing. After verificationof these distribution models, tiger locations wereused to determine the use of an area accordingto the availability and abundance of its majorprey. Details of the distribution of the preyspecies are given in Table xx. Nilgai has a verywidespread, and more or less even, distributionin all density classes. However chital was foundto have a very restricted distribution; only < 20%of the area had high chital density. Sambardistribution is also similar to that of chital; inover 50% of the area they are not found or foundat extremely low density (< 1 animal/km2).

When tiger distribution is seen in relation

to pooled animal density for these ungulates, itis observed that its distribution is closely relatedto the high prey density areas (Figure 7). Whentiger distribution was seen in relation to each ofits prey species separately it is observed thattiger distribution was not related to high nilgaidensity areas. But tiger used the high densitychital and sambar areas more than other areas,even though in terms of space these areas arehighly restricted. This indicates that tigerdistribution is related to high prey availabilitybut it is the chital and sambar availability whichis affecting the tigers’ use of habitat more thanother prey species in Panna National Park.

In a dry environment like Panna, water playsa key role and it was noticed that during thesummer months not only was prey speciesdistribution related to its availability, but tigerdistribution was also very closely related towater (Figure 33).

DiscussionHome ranges As known from other studies on the landtenure system of tiger (Sunquist, 1981, Smith,1984; Karanth and Sunquist, 2000), in Pannaboth the sexes of tiger also maintained mutuallyexclusive home ranges which can be defined asterritories. Male territories overlapped withrelatively fewer females then known from studiesconducted in Royal Chitwan Nation Park(Sunquist, 1881 and Smith, 1984). In Panna, thehome ranges of the male radio-collared tigersincluded only two three female territories,whereas in Chitwan the number included rangedfrom three to six. Territorial tigers did not sharetheir core areas with their neighbours butperipheral areas of their home ranges did overlapwith neighbouring tigers. But this overlap islimited to a small proportion of their ranges,usually less than 10% of the home range. Theiruse of core areas did not change over the years,but the shape and size of the annual home rangedid vary slightly year to year. This confirms thattiger show site fidelity once they haveestablished a territory and continue to defendand occupy the area for life. Seasonal homes in monsoon and winter offemale tigers are 22-23% smaller to the annualhome range but about 40% of the annual ranges

74

1 2 3 4 5 6

0

10

20

30

40

50

Per

cen

t

All prey abundance

Avail area Male use Female use

0 1 2 3 4 5 6 7

0.0

5.0

10.0

15.0

20.0

25.0

30.0

35.0

Per

cent

Habitat type

available Male use Female use

0

10

20

30

40

50

60

70

80

90

< 100 100 - 200 200 - 300 300 - 400 > 400

Collaredt igerCollaredt igr ess

Distance from water in metres

Fre

qu

ency

(%

)

0

10

20

30

40

50

60

70

80

90

< 100 100 - 200 200 - 300 300 - 400 > 400

Collaredt igerCollaredt igr ess

Distance from water in metres

Fre

qu

ency

(%

)

Figure 2. Habitat use by radio-collared tigers in PNP.

Figure 33. Distribution of tiger location in relation to water holes during summer months in PNP.

Figure 34. Use of space by radio-collared in relation to prey abundance in PNP.

Outsid

e NPDistur

bedOpen

hab.Thorn

scrubThorn

forestMisc.

forestMixed

forest

05

101520

2530

35

40

45

Available % use in summer % use in winter

Outsid

e NPDistur

bedOpen

hab.Thorn

scrubThorn

forestMisc.

forestMixed

forest

05

101520

2530

35

40

45

Available % use in summer % use in winter

a. Annual b. Seasonal

75

1 2 3 4

0

10

20

30

40

50

60

70

Per

cent

Chital abundance

Avail. area Male use Female use

1 2 3 4

0

1020

30

40

50

60

70

Per

cent

Sambar abundance


1 2 3 4

0

10

20

30

40

50

Per

cent

Nilgai abundance


1 2 3 4

0

10

20

30

40

50

60

70

Per

cent

Chital abundance

Avail. area Male use Female use

1 2 3 4

0

1020

30

40

50

60

70

Per

cent

Sambar abundance


1 2 3 4

0

10

20

30

40

50

Per

cent

Nilgai abundance


Figure 35. Use of space in relation to abundance of different prey speceis in PNP.

76

during summer. But a similar trend is notobserved in males; they have, on average, 40-50% smaller seasonal ranges. This indicatesthat the tigers exploit different areas accordingto the seasonal availability of the resources. Inmonsoon and winter when environmentalconditions are optimal, tigers made use of mostof the areas, including open habitats. In thesummer months open habitat are avoided andtiger distribution is limited to areas near waterholes and cool resting places. Area of intensiveuse (core area), where the tiger was located 80%of time is limited to 19% of the annual range inmales and 24% in females. An understanding ofthe characteristics of these core areas is,therefore, very important for managing an areain a way that can fulfill the minimum ecologicalrequirement of tiger. Tigers are highly mobile animals and movelong distances patrolling their territories and insearch of prey. On average males tiger covereddistances twice as large as the female tigers.The average distances travelled by tigers onlyprovides an approximate index of their movement,not the actual distances covered by the animals.No seasonal pattern is observed in distancestravelled by tigers in Panna Tiger Reserve in theway observed in seasonal home range sizes. The home ranges of tiger in Panna are largerthan those known from other tiger populationsstudied in the subcontinent (Sunquist, 1981;Smith, 1984; Karanth and Sunquist, 2000),especially that of the breeding females. Femaleterritories are mostly resource oriented(Sunquist and Sunquist, 1989). In Panna theirlarger range could well be largely due to 20to30% of unsuitable habitat, uneven distributionof prey, high human disturbance and sparsewater distribution in summer. Furthermore theheterogeneous distribution of resources overtime and space makes resource availabilityunpredictable. Larger home ranges in Panna maywell be a response to this highly spatial andtemporal unpredictable resource availability. Inorder to create more stable availability ofresources throughout the year, tigers in Pannamanage by including a larger proportion ofunproductive and less suitable habitat withintheir home ranges. As a result of this ecologicalresponse, Panna supports a smaller tiger

population currently then it could otherwise(Karanth and Nicholas, 2000). One of theimportant management aspects (other thanbiological but relevant for the conservation oftigers in Panna) related to their larger ranges isthat the tiger home ranges either extend beyondthe Park boundaries or touch the periphery. Thisexposes them to external pressures and bringsthem into direct conflict with local humanpopulations.

Tigers in Panna Tiger Reserve show distinctseasonal activity patterns. They show acrepuscular pattern with two activities bursts.This pattern is clearer during the monsoon, withleast activity during mid day. In winter most ofthe activity is seen soon after sunrise in themorning hours. The tigers remain activethroughout the day, more than they are in thenight. The least activity is noticed during thecoldest hours, between the middle of the nightto the early morning. Activity declines to itsminimum during middle of night and remainsbelow. Summer activity pattern is in sharpcontrast to winter indicating that thermo-regulatory requirement plays an important roleand governs activity patterns. The least amountof activity is seen during hottest period andactivity picks up only after sunset in the evening.

Habitat useIn Panna radio-collared tigers used denselyforested habitat more than other open habitats.Open habitats are used more only in the coolerwinter season. During summer they used denseforest habitat at the base of the steepescarpments which provide suitable cool placesto escape from the intense heat. The distributionof tigers in Panna is closely related to high preydensity areas, as has been found in other areas(Sunquist and Karanth, 1999). It is clear fromthe results of the study that tiger distributionis more closely related to two prey species ofdeer - chital and sambar - and not to nilgai, alarge bodied antelope, an otherwise ideal prey interms of size and abundance. The Dry Foresthabitat of Panna is an area occupied by twodifferent kinds of ungulate populations whichhave evolved in different environments - one inopen habitats and the other in forests. Nilgaiand chinkara represent a prey population which

77

prefer open habitats, whereas chital and sambarare forest-edge habitat dwellers. From Figure 8,it is clear that although the nilgai and chinkarapopulation occupy a much larger area, most ofit is outside the tiger distribution. Because tigerhas evolved as a specialized forest-edge predatorfollowing the cervid radiation in Asia (Sunquistand Karanth, 1999), its survival and huntingstrategies are more cued to cervids than to preyfound in open habitats. The tiger is not avoiding

Tiger-III: Food habits and predation Predators like tiger play a significant anddefinite role in the dynamics of an ecosystem.Conservation management of large carnivores isdifferent from other conservation approaches(Ginsberg, 2001). This is due to the fact thatlarge carnivores like the tiger often live in human-dominated landscapes in Asia and come intodirect conflict with humans due to theirpredation on livestock (MacDougal, 1987). Thisconflict with the human population tends to besharpest in areas where the prey population isdeclining. It is for this reason that conservationand management of tiger has always remaineda highly controversial subject forconservationists and the communities who livealongside these large predators (Joslin, 1973;Schaller, 1983; Sawarkar, 1986; McDougal,1987; Sunquist, 1981; Seidensticker et. al., 1999;Karanth, 1991; Chundawat; 1999; Gingsberg,2001). Recent studies have documented preydepletion as a major factor responsible for tigers’declines in most places (Karanth and Stith,1999). There are several factors that govern thepredation pressure on prey populations andavailability of its major prey is one of them.Another factor associated with this aspect ofabundance is the availability of alternate andsuitable prey. Apart from these factorsavailability of suitable hunting cover andcompetition play a significant role (Sunquist andSunquist, 1989). It is not always easy to gather information

on predation by large carnivores from directobservations on a regular basis. Therefore foodhabitat and predation by tiger and othercarnivores is determined from the scat analysisand kill records (Flyod et al., 19778; Sunquist,1981; Tamang, 1982; Johnsingh, 1983; Ackermanet al., 1984; Leopold and Krausman, 1986,Karanth and Sunquist, 1995, Chundawat andGogate, 1999). Scat analysis has severaladvantages over other methods (Reynolds andAebischer, 1991); it is non-invasive and scatssamples can be collected easily. But one of themajor constraints, which can affect theinterpretation of the data on the basis ofpercent occurrence, is that smaller prey itemsare usually over represented in the scat analysis(Karanth, 1993 and Karanth and Sunquist,1995). Flyod et al., 1978 found an inverserelationship between the number of fieldcollectible scats and the prey size. Based on thisrelationship Ackerman, et al., 1984, developedan equation to estimate biomass consumed perscats for cougar (Felis concolor). For estimatingthe biomass consumed by tiger, many scientists(Karanth, 1993, Karanth and Sunquist, 1995,BiswasXX) have successfully used this correctionfactor that was developed for cougar. Thisapproach is also employed in this study todetermine the food habits of tigers in Panna TigerReserve.

Results

nilgai consciously, but only a part of thepopulation of nilgai is predated - that whichoccupies forest-edge habitat. Moreover, nilgai isless likely to achieve their optimal densities inthese forest habitats. Therefore, in habitatssuch as Panna, which form over 40% of the tigerhabitat in the subcontinent, prey such as nilgaiand chinkara may be playing a limited role in theecology of tigers.

78

Science for Saving Wild Tigers in Dry Forests of the PeninsularIndia

Being a large and charismatic carnivore,the tiger has attracted considerable attentionin the conservation community the world over. Inspite of this overwhelming care, the status ofthis predator has been deteriorating rapidly.Recently, this has reached more seriousproportions than at any other time in the historyof tiger conservation in India. Three subspecies(Panthera tigris balica, P. t. virgata, and P. t.sondaica) have already become extinct, P. t.amoyensis is on the brink of extinction (Nowelland Jackson, 1996) and populations of P.t.altaica, P.t. sumatrae and P. t. corbetti are holdingon against great odds. The royal Bengal tiger, P.t. tigris, found in the Indian sub-continent, hadprovided hope for the future of the species, mainlydue to the fact that it accounted for more than60% of all the wild tiger population. Many believethat the foremost chance of saving tiger forposterity rests with India. Although India heldnearly half the world’s wild tiger population, itsfuture is precarious and is threatened bypoaching, habitat loss and loss of habitat qualitycaused by depletion of prey either due to poachingor to other forms of human disturbance (Panwar,1982; Seidensticker, 1986; Wemmer et al, 1987;Johnsingh et al. 1991; Karanth, 1993; Curry,1996; Sahgal and Thapar, 1996). A population of over 3,500 tigers in India maylook very promising, but a closer, critical look,shows all the promise to be superficial. In factthe status of the tiger in most parts of thecountry is extremely grim. Besides the problemsmentioned earlier, there are two morecharacteristics of the tiger population in India,which could lead to the future doom of the sub-species. One is the small and isolated nature ofthe tiger habitats; tigers are divided intohundreds of small populations. More than 80%of the protected areas that support tigerpopulations in India are smaller than 400 km2

and the average population size of tigers in theseprotected areas is shockingly low - less than 15individuals. Such small populations of a largecarnivore are extremely vulnerable to extinctiondue to demographic and environmental

stochasticity that operate on small populations(Gilpin and Soule, 1986; Caughley, 1994). When a tiger population of over 3,500animals is considered as one large contiguouspopulation, survival prospects look very differentfrom the reality of many scattered smallpopulations. Any modelling using the total figurewill predict a much longer survival. In fact theextinction processes are acting on each of theseisolated populations simultaneously andseparately. As a result these small and isolatedtiger populations will be lost much sooner thanone large continuous population. Tiger is not a habitat specialist: it occupiesdiverse habitats, ranging from the hot aridregions of western India, to the mangroves andhumid rainforests of tropical Asia and the Arcticclimate of Siberia (Sunquist et al., 1999). Indifferent habitat types and environments itsresponse varies; tigers achieve maximum densityin a productive and prey rich tropical climate suchas the terrai flood plains and in moist tropicalforests (Sunquist, 1981). The projection forsurvival will vary in different habitats. It is more or less an accepted fact that thetiger population in India has declined and iscontinuing to decline. Unfortunately officialfigures do not agree with expert opinion and theoverwhelming peoples’ perception that the tigerpopulation is in serious crisis due to this decline.After thirty years of tiger conservation efforts,we do not have a monitoring protocol that cantrack the changes in populations and show wherethis decline is occurring; nor is it capable of tellingus in which habitats tigers are most vulnerable.The way our official census figures are collectedgive very little indication of trends and are ofvery little use in answering such basic questions. We in India boast that if the tiger survives,it will be in India, but we prefer to ignore theprecarious status of most of the tigerpopulations in our country, which are not verydifferent from populations in the rest of theworld. Our track record in understanding theecological needs required for efficientmanagement of tigers, is extremely poor. In the

79

past thirty years of tiger conservation under theauspices of Project Tiger, which includes a strongmandate for research, there has failed to be asingle long-term study on tiger. We still have toconsult Schallers’ book, “The Deer and the Tiger”,published in 1967. Management of wildlifepopulations is a highly specialized field and inthe present scenario management can ill affordad hoc management decisions; it requiresinformed decisions. Unfortunately managementof most of the Parks in the country suffer dueto deficient scientific information on wildlifepopulations. Instead of benefiting wildlife, ad hoc,whimsical intervention, due to lack of knowledge,can and does destroy habitat and itspopulations. To address the declining status of the tigerpopulation in India it is important to haveinformation on where this decline is taking placeand what can be done to stall this decline. Sincethere is very little information available to answerthese simple questions we gathered basic dataon presence and absence of tigers in protectedhabitats (national parks and wildlifesanctuaries) from peninsular India (excluding thenorth-east states). We analysed this data todetermine where tiger was present fifty year agoand whether it is still present or absent now.This simple analysis was conducted in 1996 andprovided us with some shocking results (Table13). It also showed us where most of the declinewas taking place. It indicated that tigers havedisappeared from 70% of their original protectedhabitat in semi-arid forests and from 35% in DryDeciduous Forests. The crucial factor is thatthese two habitats together form the largest(> 46%) tiger habitat (known as Dry TropicalForest) in India. In addition, the number oflivestock per square kilometre of forest andhuman population is high in Dry Forest creatinganother biotic pressure on the ecosystem. This was an alarming result. What itindicates is that if this trend continues then alarge number of tiger populations in India will belost soon and mostly from the Dry Foresthabitats. The recent loss of tigers from SariskaTiger Reserve and Kaila Devi Wildlife Sanctuary,a part of the Ranthambhore Tiger Reserve, areexamples and vindicate our predictions made nineyears ago, highlighting the extreme vulnerability

of tigers in these habitats. Low rainfall, long, ahot dry season, extreme climatic variations andhigh anthropogenic pressures on these habitatsmake these tiger habitats sub-optimal. Any lossof breeding tigers in these small and isolatedpopulations can have disastrous consequences.Management has to be ever alert to the existingthreats to provide stability to the tigerpopulations. Therefore in order to save wild tigersimmediate attention and high priority must beattached to every tiger population found in theTropical Dry Forests of India. The other aspect we observed through theseinvestigations was that the average size of theprotected areas in these habitats is small (mean= 290 km2) and the average tiger population ineach PA is also small (< 10). This is a majorlimiting factor. If you compare the results of DryForests with other more productive systems, wefound that tigers survive in most of the TropicalMoist Forest (83%) and terai flood plains(100%)protected areas. It is clear from the datathat tigers in Dry Forest habitats survive onlyin larger PAs and this is an important factoradding to the vulnerability of the tigers in DryForest habitats. However small the tigerpopulation is in the Dry Forests, saving each andevery one of them is crucial to ensure the long-term survival of this subspecies. Conservationand management of such highly vulnerable tigerpopulations requires intensive management,especially when the PAs are small andpopulations are divided into several small units. Good conservation is always based on goodscience; unfortunately this is where wildlifemanagement of this country has lagged behindand much work is needed. Detailed scientificinformation is required on the ecology of tigers.Most of the information on tiger ecology comesfrom studies conducted in optimal habitats withhigh productivity and prey abundance and notfrom Dry Forests from where it is urgentlyrequired. That is why this study on tiger in PannaTiger Reserve is important and was designed tofind out the ecological requirements needed tomaintain a demographically viable population andidentify factors responsible for the decline oftiger population in the Dry Tropical Forests. Untilthis study, almost nothing was known from thishabitat, which is a typical and large tiger habitat

80

17001170236(70%)

350(30%)

270(n=26)

Semi-arid

563881230(35%)

400(65%)

316(n=39)

Dry deciduous

3401600<100(10%)

400(90%)

342(n=8)

Rain forest

269573180(17%)

774(83%)

660(n=63)

Moist deciduous

6001800nil418(100%)

418(n=12)

Terrai flood plains

Tiger lost

Tiger present

All PAs

Cattle /km2 of forest

Human /km2 of forest

Average Protected Area size in km2

Habitat types

All= average size of all protected tiger habitatTiger present= average size of protected tiger habitat where tiger still occur;Tiger lost= average size of the protected tiger habitats from where tiger have been lost;n= number of protected areas sampled;Figure in bracket= percent of protected areas;Humana and cattle/km2 of forest= rural human and cattle population per km2 of forest

17001170236(70%)

350(30%)

270(n=26)

Semi-arid

563881230(35%)

400(65%)

316(n=39)

Dry deciduous

3401600<100(10%)

400(90%)

342(n=8)

Rain forest

269573180(17%)

774(83%)

660(n=63)

Moist deciduous

6001800nil418(100%)

418(n=12)

Terrai flood plains

Tiger lost

Tiger present

All PAs




Habitat types

17001170236(70%)

350(30%)

270(n=26)

Semi-arid

563881230(35%)

400(65%)

316(n=39)

Dry deciduous

3401600<100(10%)

400(90%)

342(n=8)

Rain forest

269573180(17%)

774(83%)

660(n=63)

Moist deciduous

6001800nil418(100%)

418(n=12)

Terrai flood plains

Tiger lost

Tiger present

All PAs




Habitat types

All= average size of all protected tiger habitatTiger present= average size of protected tiger habitat where tiger still occur;Tiger lost= average size of the protected tiger habitats from where tiger have been lost;n= number of protected areas sampled;Figure in bracket= percent of protected areas;Humana and cattle/km2 of forest= rural human and cattle population per km2 of forest

of peninsular India but where the tigers are mostvulnerable. Findings from this research reveal howresilient the tigers are but also warns howinsecure populations like Panna tigers can be ifmanagement is lax and not attentive to existingthreats. Tigers are specialized hunters and theirresponses vary according to variable ecologicalconditions. It is, therefore, important to studytheir behavioural responses because it can helpus understand why tigers are so vulnerable inits largest habitat, the Dry Tropical Forest. Forexample in a prey depleted area, tigers may bekilling livestock more often than expected,bringing the population into direct conflict withhumans. The response of the tiger populationcould be low survival rates and poor reproductivesuccess and the end result would be a highlyvulnerable tiger population. To understand tiger ecology it is importantthat we gather very reliable information. Thisrequires unbiased information on its ecology; toachieve this, it is essential to locate tigers forobservations at will. But tigers are difficult tolocate and observe and direct observations are

limited in the wild. A few opportunisticobservations from direct sightings, andassumptions based on these limitedobservations, could be highly biased andtherefore misleading. Radio-telemetry is,therefore, an important tool for gatheringinformation in an unbiased manner and in muchgreater detail. An unexpected spin off of intensiveand constant monitoring of radio-collared tigersis that it can reduce substantially any humancaused mortality and provide security in a wayno other monitoring can. One of the major findings of this study inPanna is that the home ranges of tigers are muchlarger than known from other, more productive,habitats of the sub-continent. Male tigers onaverage ranged over 210 km2 and females 58km2. These ranges are two to three times largerthan those estimated from Chitwan andNagarahole National Park. This is a significantfinding. Not because it is new information butbecause it has serious conservation implications,especially in a Dry Forest habitat. Larger homeranges mean that tiger densities in Panna arelow and fewer breeding territories could be

Table 1. Assesment of tiger vulnerability in different bio-unit/ forest types of Penninsular India.

81

70.7446.43552.7average

37.5720.0375.62Panna NP


73.9144.1356.13Nagarahole. NP


91.0980.8118.76Kanha NP

81.6658.9141.08Pench NP

NumberMedium Prey

BiomassMedium prey

BiomassLarge prey

ProtectedArea

70.7446.43552.7average

37.5720.0375.62Panna NP


73.9144.1356.13Nagarahole. NP


91.0980.8118.76Kanha NP

81.6658.9141.08Pench NP

NumberMedium Prey

BiomassMedium prey

BiomassLarge prey

ProtectedArea

accommodated within the protected area. Thisis an important finding because it is anestablished fact from other studies thatpopulations with larger home ranges are highlyvulnerable to local extinction. Therefore, if largerranges are a typical characteristic of Dry Foresthabitat, it can be safely stated that tigerpopulations of Dry Forests are highly vulnerable.However, in sharp contrast to the need for largerareas, PAs of Dry Forest are small (mean=316km2). They can support fewer breedingindividuals and these small populations, even inthe best of situations, will find it hard to survive.This is one of the major reasons why tigers indry and semi-arid areas are disappearing at afaster rate. Tiger avoids open forest in Panna and it isestimated that approximately 30% of thehabitat is naturally unsuitable for tigers in theReserve because it is avoided by them. Thisfurther adds complexity for the management inunderstanding the minimum ecologicalrequirements of tigers. Large home ranges oftigers in Panna could be one of the responses tothis ecological condition. As a result fewerbreeding territories could be accommodated inPanna. During this study we identified a maximumof only seven breeding female territories in theReserve, south of the river Ken. This accountsfor only two breeding units belonging to twodominant males with three to four femaleterritories each. This is an extremely smallbreeding population and unlikely to ensure long-

term survival of the population. Another aspect which adds to vulnerabilityof the tiger population, associated with the largehome ranges is that the territories of all thebreeding tigers monitored by the research team,extend either up to the periphery of the Reserveor beyond its boundaries into the tiger habitatsavailable outside (see map page…). This is whereconflict rules; tiger here predate mainly onlivestock and come in direct conflict with the localhuman population, exposing themselves tohuman caused mortalities. Since most of theprotected areas in Dry Forest habitats aresmaller then Panna, it is likely that a similarsituation exists in most of them. Thus none ofthese breeding tigers are fully protected withinthe Reserve. Management can provide all theprotection within its boundaries but when itsbreeding tigers venture outside its managementjurisdiction they are unsafe. It clear from themonitoring of tigers in Panna how unsafe thesebreeding individuals are. Out of the known elevenbreeding tigers monitored by the research team,Panna Tiger Reserve has lost eight of them. Thisis a significant loss of breeding tigers. Such highlosses and even turnover of breeding populationdoes not augur well for the tigers’ long-termviability. When this is true for breeding individuals,it is difficult to provide stability to the breedingpopulation. To achieve maximum reproductivesuccess and save wild tigers, stability of thebreeding population is an essential requirement.This is an important issue in saving the tiger in

Table . Percent biomass and number of tiger prey in different protected areas.

82

its largest habitat and one that needs to beurgently addressed by wildlife managers.Immediate action would be to include, whereverpossible, all the adjoining tiger habitats withinthe PA management as buffer, with immediateeffect. This can be done without a change in theirlegal status; what is important, is that thesecrucial breeding areas must come under onemanagement unit to ensure stability of thebreeding population. From the studies conducted in Panna, it isclear that survival of a tiger population is notdependent on the total number of tigers it hasbut on the size of the breeding population itsupports. From the study we learned that notall tigers in a population are territorial - ratherit consists of territorial (breeding) and non-territorial (transient sub-adult cubs and fewernon breeding adults) individuals. The territorialpopulation usually remains constant (in idealcondition) and it is the non-territorial populationthat fluctuates. Any instability in the breedingpopulation can be disastrous. In a typical tigerpopulation there would be a larger proportion ofnon-territorial and a smaller proportion ofterritorial animals. The viability of the populationis dependent on how large or small this breedingproportion is and its reproductive success. Forexample, if you have a population of hundredtigers in an area but only 10% of them arebreeding, the population may not be viable in thelong term in spite of it being a large population.This is because it mainly consists of a non-territorial population, which fluctuates andcontributes little to the viability of thepopulation. Therefore, to save a tiger populationin the wild, management needs to maintain anappropriate proportion of territorial populationwithin the protected area and provide thispopulation with stability and an environmentsuitable for its successful breeding. Preliminaryanalysis of our data indicates that a minimumof nine female (breeding) territories would keepa Panna population of 35 tigers viable for a longperiod. Finding space for these nine breedingterritories is not easy and it becomes moredifficult when other ecological factors need tobe considered. Not all the territories will be equalin habitat quality and fulfil minimum requirementof a breeding territory. This can reduce the over

all reproductive success of the population. Tocompensate for this, additional breedingterritories must be identified, included andprotected to ensure successful breeding in thearea. These are difficult goals to achieve in areaswhere human populations dominate the tigerhabitat. Therefore, for a stable and viable tigerpopulation, an inviolate area large enough tosupport a breeding population capable ofsustaining a viable tiger population is essentialand must not be compromised if we wish toensure that tiger populations survive for decadesto come. Our efforts to save wild tigers are helped bythe fact that tiger is a resilient species. With alittle support it can bounce back, if its minimumecological requirements are satisfied and human-caused mortalities are kept to a minimum. Thisis what was witnessed in Panna between 1996and early 2002. We documented one of the finestrecoveries of a tiger population from 2-3 tigersper 100 km2 in 1996, when the project started,to 7 tigers to every 100 km2 in 2002. We havemonitored tigress-123 and her family for overseven years. During this period we saw 26 cubsproduced by her and her children. We havewitnessed three generations (grandchildren) oftigers, their survival, breeding and land tenuresystems. Out of these 26 cubs, 20 survived toadulthood and dispersed. Such a high survivalrate is a remarkable achievement for themanagement and indicates the reproductivepotential of tigers. Long term monitoring of thisfamily unit in one neighbourhood provided us withinsights that can help management plan astrategy to save tigers based on objectiveinformation now available to them from thisstudy. What characterizes a breeding habitat?What are the ecological requirements for tigersto breed successfully? Such basic informationis important for management to know. This studydocuments the ecological requirements of tigerin detail for the benefit of the management. Toachieve an optimal reproductive success it isimportant that all breeding territories must fulfilcertain minimum ecological requirements. Infailing to do so, reproductive success of thepopulation and ultimately its survival, is affected.In Panna, when monitoring adjoining female

83

territories over the study period, it was observedthat the tigresses’ responses differ with thevariable ecological conditions that exist in theseterritories. One of the female’s territories(female-123) has an even distribution ofresources, such as high uniform distribution ofprey density, water, cover and no disturbance.The result was a high reproductive success andthe tigress was able to raise most of her cubs(almost 90% survival of 11 cubs in four litters3+2+3+3) successfully and survived throughoutthe study period. However, in neighbouringterritories, which are characterized by unevenresource distribution of water, cover and preydensities, higher livestock population andincluded larger disturbed habitats, the tigresseshad only limited (<40 %) reproductive successesin raising the cubs to adulthood. Moreover, noneof the breeding females survived the entire studyperiod and died within five to seven years. Thisvariability in the quality of the breeding habitatin the system affects the over all reproductivesuccess and viability of the tiger population. It is estimated from the study that tigersin Panna kill more than 6 cattle annually, whichtranslates to over 30% of the prey biomassconsumed and approximately 15% of the numberof prey killed. Observation of kill data of radio-collared tigers show that contribution by cattleto the tiger’s diet could be as high as 40% andeven higher for male tigers. In order to maximiseits reproductive success a male tiger covers largeareas to encompass several female territories.Its large range includes a greater proportion ofdisturbed habitats. Because strategy of a maletiger is mainly mate oriented, he is constantlyon the move and patrols his territory extensivelyto fight off any challenger. This occupation keepshim very busy and forces him to move largedistances. A male tiger, therefore, more oftenencounters and takes advantage of this easyprey. Such persistent killing of livestock by maletigers, as observed in Panna, brings them intodirect conflict with the local people. The maletiger plays a very crucial role in tiger biology. Maletigers not only bring fresh genes into thepopulation and contribute three to four timesmore cubs then a female but, most importantly,he also ensures cubs survival. A constantchangeover of males as seen in Panna in the later

part of the study between 2002 and 2005,adversely affects the reproductive success ofthe population. If human caused mortality onmales is high, its effect on the population canbe catastrophic and will seriously jeopardize itssurvival.

The females’ strategy on the other handis resource oriented because they spend mostof their time raising cubs. They occupy areaswhere food and other resources are in abundance.We now know how much food, what kind of preyand where and when it is needed, through thisstudy. The tiger’s food can be classified aspreferred and principal. ‘Preferred’ is food thatis eaten more often when given a choice. ‘Principal’food is the staple diet – food that is eatenfrequently and forms the bulk of the intake. Thestudy suggests that at least one of the majorprey species of tiger should always be present inabundance to be available as the principal food.When prey availability and composition of Pannais compared with other high tiger density areas,it is obvious that this is not the case here. Threemajor ungulate prey species - nilgai, sambar andchital - form the bulk of the ungulate biomass inPanna. The nilgai’s preference for open habitatsprecludes it from substituting as the principalprey for tigers in dry forest habitat. In Panna,tigers do not kill nilgai as much as one mightexpect from its availability. These large antelopes,well suited for open habitat, are predated onlywhen frequenting forested habitats alsofrequented by tigers. Tigers do not purposelyavoid nilgai but they do not frequent nilgaihabitat often. Wild ungulate biomass from openand disturbed habitats does not, therefore,necessarily translate into high prey availabilityfor tigers. Besides, nilgai, like sambar, cannotbecome the tigers’ principal food as neither canachieve the densities that chital can. Thepotential principal food for tigers in Panna istherefore chital, like it is in most of the tigerhabitats of the sub-continent. On average theprincipal food, or chital, contributes about 46%of the total prey biomass and over 70% in termsof number animals in high tiger density areassuch as Kanha, Nagarahole and Kaziranga.Whereas in Panna, chital are limited in numberand also restricted in distribution, itscontribution is only 20% of available prey biomass

84

and 37% in term of number. Other large prey -sambar and nilgai - together on averagecontribute over 70% to the prey biomass but cannever achieve the abundance frequently reachedby chital in the sub-continent. Chital and othersimilar sized prey such as wild pig and hog deercan attain higher densities and their superabundance makes it possible for a tiger to getfood at regular intervals. The collared tigressraising three one year old cubs kills, on average,a medium to large bodied prey once every six days.Low availability of medium size prey iscompensated by increased intake of smaller prey,such as langur and four horn antelope, in Panna.The role of medium size prey as principal food inthe ecology of the tiger is an important factoraffecting the tiger’s responses (such as tigerdensities, land tenure, distribution andreproductive success) significantly more thanother prey species. A wider availability of principalfood with increased abundance will have a positiveeffect on restoring tiger ecology in Dry Forests. In Dry Forests, at the end of the growingseason when most of the pastures are grazed,pressures intensify on the intact plant biomassof the Protected Areas. In these areas, livestockpopulations are dependent on forest resourcesfor most of the year. The cumulative effect ofhigher human pressure, altered ecologicalconditions, unpredictable resource distribution,high conflict, low prey availability and smallerbreeding units makes tiger populations in theseareas more susceptible to the extinctionprocess; a slight perturbation will have apronounced effect on them. One must attend to these problems at theearliest, otherwise India could lose a large partof its tiger population very soon. Increasing thesize of PAs is not always possible, therefore analternative management strategy is required tosave these populations. Wildlife populations andtheir habitats are dynamic systems and requireconstant monitoring and intervention to protectits characteristic (biodiversity) andfunctionality. Wildlife management in Indiarequires scientific information to direct themanagement actions on the right path, help planmanagement strategies and save our naturalsystem. We need to move beyond the oldphilosophy of protecting an area and leaving

nature to take its own course. Given the currentstatus of our wildlife, we can ill afford to sit backand let the situation be. We do not have theluxury of large areas free from humandisturbance where nature can play its true role,unimpeded.

After the great tiger population recoverysuccess story, the sudden loss of tigers in Pannaproves just how inadequate our monitoringsystems are and limited our managementsystem is. What was observed in Panna after2002 is what makes tiger populations sovulnerable to extinction in Dry Forests. In 2002,the tiger population was estimated to be 29tigers (including cubs) in 410 km2 of the sampledarea using the photographic capture-recapturetechnique. This included seven breeding females,all with a litter of two cubs each except one whohad a litter of three cubs. This accounted forfifteen cubs of different age groups but most ofthem over one year old. These female territorieswere dominated by two territorial breeding males.In addition to this there was evidence of four tofive other adult non-territorial tigers thatincluded a male cub of female-123 from herprevious litter. But once it was believed that thetiger population was safe and managementbecame lax, mortalities, mainly human caused,struck at regular intervals and Panna TigerReserve lost most of its breeding population. Outof the eleven breeding tigers known to theresearch team, nine of them disappeared. Almostno cubs were recorded by the censusesconducted by the staff and the research team.Most of the cubs and sub-adult tigersdisappeared too; survival rates declined sharply.This loss of tigers in Panna highlights why tigerare vulnerable in Dry Forest. Losses of tigers inSariska and Kaila Devi exemplify the past trendwe have noticed from our presence/absenceanalysis of tiger populations. It is now very clearfrom the examples of these losses from Sariska,Kaila Devi and Panna that if we are going to losetigers in India, it will be first from Dry Foresthabitats. Less than 4% of the country comes underthe protected area network and 80% of theProtected Areas are less then 400 km2 in size.They exist in a populous country with manyhuman demands on the natural resources. In

85

such circumstances management has to workintensively and in so doing cannot afford to relyon guesses based on individual opinions. We needhard facts and information in order to developcreative solutions to the problems faced by PAmanagement. Such serious data and informationcan only come from scientific studies like thisone. This is where wildlife science plays a crucialrole in wildlife conservation. It cannot save tigersbut can guide management towards anappropriate path. We need constant monitoringof our wild heritage because managementproblems will never disappear, they will changeover time and new ones will appear. Scientificknowledge of the situation is the only way forwardto take informed actions to mitigate each newset of problems. Otherwise, uninformed actionscan create more problems than they solve. Overthe past three decades the role and importanceof science in wildlife conservation has beenhighlighted and discussed in every national andinternational forum yet very little progress has

been made. Due to lack of progress we still haveto refer to the DEER AND the TIGER and THEBOOK of INDIAN ANIMALS or, when more detailedinformation on some common animals such assambar, nilgai and chital is sought, we have torefer to those written on introduced populationsin Texas, USA. We need to correct this, hopefullyin the coming decades we will be able to changethe status of wildlife science in India so thatwildlife management will be equipped withappropriate and adequate information. In theabsence of scientific knowledge wildlifemanagement will remain in its infancy. Betterunderstanding of the problems and innovativeactions to protect these highly dynamicbiological systems will help management evolveinto a more mature and effective system. Onlyin this way can we take conservation forward intothis new 21st century so that tigers and all theother diverse array of the natural world in Indiamay exist for generations to come.

86

Literature CitedAnderson, D. R., Burnham, K. P., Lubow, B. C., Thomas, L., Stephen Corn P., Mrdica, P. A. andMarlow, R. W. (2001). Field trials of line transect methods applied to estimation of desert tortoiseabundance. J. Wildl. Manage. 65:583-597.

Anonymous, 1993. Proceedings and resolutions. International Symposium on the Tiger, NewDelhi. Project Tiger, Ministry of Environment and Forests, Government of India.

Biswas, S. & Sanker, K. (2002). Prey abundance and food habit of tiger (Panthera tigris tigris) inPench National Park, Madhya Pradesh, India. J. Zool. Lond. 256:411-420).

Borchers, D.L., Anderson, D.R., Burnham, K.P., Hedley, S.L., Pollard, J.H. and Bishop,J. R. B. 2003. Distance 4.1. release-2. Research Unit for Wildlife Population Assessment,University of St. Andrews, UK. http://www.ruwpa.st-and.ac.uk/distance/

Buckland, S. T., Anderson, D. R., Burnham, K. P. & Laake, J. L. (1993). Distance sampling:estimating abundance of biological populations. London: Chapman & Hall.

Buckland, S. T., Anderson, D. R., Burnham, K. P., Laake, J. L., Brochers, D. L. and Thomas, L.(2001). Introduction to distance sampling: Estimating abundance of biological populations.Oxford: Oxford University Press.

Burnham, K P and D R Anderson, 1984. The need for distance data in transect count. Journal ofWildlife Management. 48:1248-1254.

Burnham, K P, D R Anderson, J L Laake, 1980. Estimation of density from line transect samplingof biological population. Wildlife Monographs. 72. The Wildlife Society, Washington DC. 202pp.

Burnham, K.P. & Overton, W.S. (1978). Estimation of the size of a closed population whencapture probabilities vary among animals. Biometrika 65: 625-633.

Burt, W. H. 1943. Territoriality and home range concepts as applied to mammals. Journal ofMammalogy. 24: 346-352.

Byers, C. R., Steinhorst, R. K. 1984. Clarification fo a technique for analysis of utilization-availability data. J. Wildl. Manage. 48(3): 1050-1053.

Carbone, C., Christie, S., Conforti, K., Coulson, T., Franklin, N., Ginsberg, J.R., Griffiths, M.,Holden, J., Kawanishi, K., Kinnaird, M., Laidlaw, R., Lynam, A., MacDonald, D.W., Martyr, D.,McDougal, C., Nath, L., O’Brien, T., Seidensticker, J., Smith, D., Sunquist, M., Tilson, R. & WanShahruddin, W.N. (2001). The use of photographic rates to estimate densities of tigers and othercryptic mammals. Anim. Conserv. 4: 75-79.

Champion H. G. and Seth, S. K. 1968. The revised survey of the forest types of India. Manager ofPublications. Govt. of India. New Delhi. 404 pp.

Chaudhary, P.K. 1996. Economic Dependence of Enclaved and Surrounding Villages on PannaNational Park - A Search for an Alternative. Consultancy Project Report under GOI-UNDPProject.

87

Chawdhry, P.K. 1997. Management Plan for Panna National Park (Period 1997-98 to 2006-07) -A draft report. Forest Department, Government of M.P.

Chellam, R. 1993. Ecology of the Asiatic Lion (Panthera leo persica). PhD. thesis, SaurashtraUniversity, Rajkot, India. Pp. 170.

Chundawat, 2001. Tiger conservation Dry Tropical Forests of India. Technical report: 2000-2001.

Chundawat, R. S. & Gogate, N. (2001). Saving tigers in Dry Tropical habitats. In Saving wildtigers. Thapar, V. (Ed). Permanent Black, Delhi. 385-394.

Chundawat, R. S., N. Gogate and A.J.T. Johnsingh, 1999. Tiger of Panna: Preliminary resultsfrom an Indian Tropical Dry Forest. In Riding the tiger: Tiger conservation in human-dominatedlandscape. Seidensticker, J., Christie, S. 7 P. Jackson (Eds). Cambridge: Cambridge UniversityPress.

Debroy, S. (1996). Halting the decline of the tiger. J. Wildl. Res. 1(2): 224-228.

Dixon, K. R. and J. A. Chapman, 1980. Harmonic mean measure of animal activity areas.Ecology. 61(5): 1040-1044.

Dobson,F.S. 1982. Competition for mates and predominant juvenile male dispersal in mammals.Anima. Behav. 30:1183-1192.

Eberhardt, L.L. 1968. A preliminary apprasial of line transect. J. Wildl. Manage. 32: 82-88.

Eisenberg, J F and J Seidensticker, 1976. Ungulates in southern Asia: a consideration of biomassestimates for selected habitats. Biological Conservation 10:293-308.

Eisenberg, J. F. & Lockhart, H. (1972). An ecological reconnaissance of Wilpattu National Park,Ceylon. Smithson. Contribution. Zool. 1001:1-118.

Eisenberg, J. F. 1981. The mammalian radiations: an analysis of trends in evolutions, adaptation,and behaviour. The University of Chicago Press. Chicago.

Ellner, S. and L. A. Real. 1989. Optimal foraging models for stochastic environments: are wemissing the point? Comments. Theoretical Biology. 1: 129-158.

Garshelis, D. L. 2000. Delusion in habitat evaluation: measuring use, selection and importance.In Research techniques in animal ecology: controversies and consequences, eds. Boitani, L. andFuller, T. K. Columbia University Press. Pp 111-164.

Gates, C.E. 1980. Linetran, a general computer program for analysing line transect data. J. Wildl.Manage..55:393-396.

Gee. E. P., 1964. The wildlife of India. London.

Hall, L. S. Krausman, P. R. and Morrison, M. L., 1997. The habitat concept and a plea forstandard terminology. Wildlife Society Bulletin. 25:173-182.

88

Jathanna, D., Karanth, K. U., Johnsingh, A. J. T. (2003). Estimation of large herbivore densitiesin the tropical forets of southern India using distance sampling. J. Zool., London. 261: 285-290.

Jennrich, R. and E. Turner, 1969. Measurement of non circular home range. Journal ofTheoretical Biology. 22:227-237.

Johnsingh, A. J. T. (1983). Large mammalian prey-predator in Bandipur. J. Bombay Nat. HistorySoc. 80:1-57.

Johnsingh, A.J.T., H.S. Panwar and W.A. Rodgers. 1991. Ecology and conservation of largefelids in India. In Wildlife conservation: present trends and perspectives for the 21st century.Proceeding of the International Symposium on Wildlife Conservation in Tsukuba and Yokohama,Japan.

Johnsingh, AJT, 1983. Large mammalian prey-predators in Bandipur. Journal of Bombay NaturalHistory Society. 80, 1-57.

Johnson, D. H., 1980. The comparison of usage and availability measurement for evaluatingresource preference. Ecology. 61:65-71.

Johnson, T. H. 1984. Habitat and social organization roe deer (Capreolus capreolus). Ph. D.Thesis. University of Southampton.

Karanth K. U. 1993. Predator-prey relationships among the large mammals of NagarholeNational Park, India. Ph.D. thesis, Mangalore University.

Karanth, K and M E Sunquist, 1992. Population structure, density and biomass of largeherbivores in the tropical forests of Nagarahole, India. Journal of Tropical Ecology 8:21-35.

Karanth, K and M E Sunquist, 1995. Prey selection by tiger, leopard and dhole in tropical forests.Journal of Animal Ecology 64, 439-450.

Karanth, K. U. & Chundawat, R. S. (2002). Ecology of the tiger: Implications for populationmonitoring. In Monitoring Tigers and their Prey: A manual for researchers, managers andconservationists in Tropical Asia: 9-21. Karanth, K. U. & Nichols, J.D. (Eds). Bangalore: Centrefor Wildlife Studies.

Karanth, K. U. & M. E. Sunquist, 2000. Behavioural correlates of predation by tiger (Pantheratigris), leopard (Panthera pardus) and dhole (Cuon alpinus) in Nagarahole, India. J. Zool. Lond.250, 255-265.

Karanth, K. U. & Nichols, J. (1998). Estimation of tiger densities in India using photographiccaptures and recaptures. Ecology. 79(8): 2852-2862.

Karanth, K. U. & Nichols, J. D (eds), 2002. Monitoring tigers and their prey: a manual forresearchers, managers and conservationists in Tropical Asia. Natraj Publication. Pp. 111-120

Karanth, K. U. & Sunquist, M. E. (2000). Behavioural correlates of predation by tiger (Pantheratigris), leopard (Panthera pardus) and dhole (Cuon alpinus) in Nagarahole, India. J. Zool., Lond.250: 255-265.

89

Karanth, K. U. (1991). Ecology and management of the tiger in tropical Asia. In Wildlifeconservation: present trends and perspectives for the 21st century. (Eds) N. Maruyama, B. Bobek,Y. Ono, W. Reglin, L. Bartos & R. Ratcliffe, pp 156-159. Japan Wildlife Research Centre, Tokyo.

Karanth, K. U. (1995). Estimating tiger Panthera tigris populations from camera-trap data usingcapture-recapture models. Biol. Conserv. 71: 333-338.

Karanth, K. U. (1999). Counting tigers with confidence. In Riding the tiger: tiger conservation inhuman-dominated landscapes: 350-353. Seidensticker, J., Christie, S. & Jackson, P. (Eds).Cambridge: Cambridge University Press.

Karanth, K. U. (2003). Tiger ecology and conservation in the Indian subcontinent. J. Bombay.Nat. Hist. Soc., (In press).

Karanth, K. U. and B. M Stith, 1999. Prey depletion as a critical determinant of tiger populationviability. In Riding the tiger: Tiger conservation in human-dominated landscape. Seidensticker,J., Christie, S. 7 P. Jackson (Eds). Cambridge: Cambridge University Press.

Karanth, K. U., J. D. Nichols, N. S. Kumar, W. A. Link and J. E. Hines, 2004. Tigers and theirprey: predicting carnivore densities from prey abundance. PNAS. 101(14): 4854-4858.

Karanth, K. U., Kumar, N. S. & Nichols, J. D. (2002). Field surveys: Estimating absolutedensities of tigers using capture-recapture sampling. In Monitoring Tigers and their Prey: Amanual for researchers, managers and conservationists in Tropical Asia: 139-152.

Karanth, K. U. & Nichols, J.D. (Eds). Bangalore: Centre for Wildlife Studies.Karanth, K. U., Nichols, J. D., Seidensticker, J., Dinerstein, E., Smith, J.L.D., McDougal, C.,Johnsingh, A.J.T., Chundawat, R.S. & Thapar, V. (2003). Science deficiency in conservationpractice: The monitoring of tiger populations in India. Anim. Conserv. 6: 141-146.

Karanth, K.U. 1987. Tigers in India: A critical review of field censuses. In Tigers of the World:The Biology, Biopolitics, Management and Conservation of an Endangered Species, eds. R.L.Tilson and U.S. Seal, pp. 118-33. Park Ridge, New Jersey, Noyes Publications.

Kawanishi, K. (2002). Population status of tigers in a primary rainforest of peninsular Malaysia.Ph.D. dissertation. Gainesville: University of Florida.

Khan, J.A., R. Chellam, W.A. Rodgers and A.J.T. Johnsingh, 1996. Ungulate densities andbiomass in the tropical dry deciduous forests of Gir, Gujarat, India. J. Trop. Ecol. 12: 149-162.

Khan, J. A. 1990. Gir Lion project: ungulate-habitat ecology in Gir. Wildlife Institute of IndiaDehra Dun. 214pp.

Khan, J. A. 1997. Estimation of ungulates densities by line transects method in Gir forest, India.Tropical Ecology 38(1):65-72.

Khan, J. A. (1996). Factors governing the habitat occupancy of ungulates in Gir Lion Sanctuary,Gujarat, India. Int. J. Ecology. Environ. Sci. 22:73.83.

90

Krebs, C. J. 1994. Ecology: the experimental analysis of distribution and abundance.HarperCollins College Publishers. New York.

Kumar, N. Samba, 2000. Ungulate density and biomass in the tropical semi-arid forest ofRanthambhore, India. M.Sc. dissertation. Pondichery University, Pondichery.

Laake, J. L., Buckland, S. T., Anderson, D. R. & Burnham, K. P. (1998). Distance: User’s guide.Fort Collins, CO: Colorado State University.

Laake, J.L., K.P. Burnham, D.R. Anderson. 1979. User’s manual for program TRANSECT. UtahState University Press, Logan. 26pp

Laake, J.L., S.T. Buckland, D.R. Anderson and K.P. Burnham. 1994. Distance: User’s guide.Colorado State University. Fort Collins.Lancia, R.A.,J.D. Nichols and Pollock, 1994. Estimating the number of animals in wildlifepopulations. In Research and Management Techniques for Wildlife and Habitats (ed.)T.A.Bookhout. The Wildlife Society, Bethsda, Maryland. 215-253.

Madhya Pradesh Gazette, Part I, published on 8th February 1985.

Marques, F. F. C., Buckland, S. T., Goffin, D., Dixon, C. E., Borchers, D. L., Mayle, B. A. andPeace, A. J. (2001). Estimating deer abundance from line transect surveys of dung: sika deer insouthern Scotland. Journal of applied Ecology.

Mathai, M.V. 1999. Habitat occupancy by tiger prey species across anthropogenic disturbanceregimes in Panna National Park, Madhya Pradesh. M.Sc. Dissertation thesis, SaurashtraUniversity, Rajkot, India. Pp. 61.

Mathur, V. B. 1991. The ecological interaction between habitat composition, habitat quality andabundance of some wild ungulate in India. Ph. D. Thesis. University of Oxford.

May, R.M. 1975. Pattern of species abundance and diversity. In Ecology and Evolution ofCommunities (ed.) M.L. Cody and J.M. Diamond. Harvard University Press.

McDougal, C. 1977. The Face of the Tiger. London, Rivington Books.

McDougal, C.W. 1975. The face of the tiger. Revington Books and Andre Deutsch. 180pp

Mech, L.D. 1983. Handbook of animal radio tracking. University of Minnesota Press.

Miquelle, D.G., Smirnov, E.N., Quigley, H.G., Hornocker, M.G., Nikolaev, I.G. and

Matyushkin, E.N. 1996. Food habits of Amur tigers in Silkhote-Alin Zapovednik and the RussianFar East, and implications for conservation. Journal of Wildlife Research, I(2), 138-47.

Mishra, H., 1982. The ecology and behaviour of chital (Axis axis) in the Royal Chitwan NationalPark, Nepal: with comparative studies of hog deer, sambar and barking deer. Ph. D. Thesis,University of Edinburgh.

91

Mueller-Dombois, D. and E. Ellenberg. 1974. Aims and methods of vegetation ecology. Johnwiley &Sons. New York.

Mukherjee, S., S.P. Goya.l and R. Chellam. 1994. Refined techniques for the analysis of Asiaticlion (Panthera leo persica) scats. Acta.Theriologica. 39(4): 425-430.

Neu, C. W., Byers, C. R., Peek, J. M.1974. A technique for analysis of utilization-availabilitydata. J. Wildl. Manage. 38(3): 541-545.

Nichols, J. D. & Karanth, K. U. (2002). Statistical concepts: Estimating absolute densities ofTigers using Capture-recapture sampling. In Monitoring Tigers and their Prey: A manual forresearchers, managers and conservationists in Tropical Asia: 121-137.

Karanth, K. U. & Nichols, J.D. (Eds). Bangalore: Centre for Wildlife Studies.Nichols, J.D. (1992). Capture-recapture models: Using marked animals to study populationdynamics. BioScience 42:94-102

Nowell, K. & P. Jackson, 1996. Wild Cats: Status Survey and Conservation Action Plan. Gland,Switzerland, IUCN.

O’Brien, T.G., Kinnaird, M. F. & Wibisono, H.T. (2003). Crouching tigers, hidden prey:Sumatran tigers and prey populations in a tropical forest landscape. Anim. Conserv. 6: 31-139.

Otis, D.L., Burnham, K.P., White, G.C. & Anderson, D.R. (1978). Statistical inference fromcapture data on closed animal populations. Wildl. Monogr. 62:1-135.

Pabla, H.S. 1984. Panna National Park - Prospects and Problems, M.P. Forest Department.

Panwar, H. (1987 Project tiger: the reserve, the tiger and their future. In Tigers of the world: thebiology, bio politics, management and conservation of an endangered species (Eds) R.L. Tilsonand U.S. Seal. 110-117. Noyes Publication, Park Ridge, New Jersey.

Panwar, H. S. 1979. Population dynamics and land tenures of tigers in Kanha National Park.Proceedings of International Symposium on Tigers, New Delhi.

Panwar,H.S. 1982. What to do when you have succeeded: Project Tiger ten years later. InNational Parks, Conservation and Development (eds.) Mcneely, J.A. and K.R.Miller).Smithsonian Press, Washington D.C.

Plumtre, A. J. (2000). Monitoring mammal populations with line transect techniques in Africanforests. Journal of Applied Ecology 37:356-368.

Powell, R. A. 2000. Animal home ranges and territories and home range estimators. In Researchtechniques in animal ecology: controversies and consequences, eds. Boitani, L. and Fuller, T. K.Columbia University Press. 65-110pp.

Prater, S H, 1971. The book of Indian animals. Bombay Natural History Society, Bombay.

Prater, S. H. 1967 The book of Indian Animals. Bombay Natural History Society. Bombay.

92

Pulliam, H.R. 1988. Sources, sinks and population regulation. Am.Nat. 132:757-785.

Pulliam, H.R. and B.J. Danielson. 1991. Sources, sinks and habitat selection: a Landscapeperspective on population dynamics. Am. Nat. 137:S50-S66.

Putman, R. 1988. The natural history of deer. Cornell University Press, New York.

Pyke G. H., H. R. Pulliam and E. L. Charnov. 1977. Optimal foraging: a selective review oftheory and tests. Quarterly Review of Biology. 52:137-154.

Pyke, G. H. 1984. Optimal foraging theory: a critical view. Annual Review of Ecology andSystematics. 15:523-575.

Rabinowitz, A. 1989. The density and behaviour of large cats in a dry tropical forest mosaic inHuai Kha Khaeng Wildlife Sanctuary, Thailand. Natural History Bulletin of the Siam Society, 37(2), 235-51.

Rexstad, E. & Burnham, K.P. (1991). User’s guide for interactive program CAPTURE.Abundance estimation of closed animal populations. Fort Collins: Colorado State University.

Rodgers, W.A. and H.S. Panwar. 1988. Planning a wildlife protected area network in India. Vol. I& II, Wildlife Institute of India.

Sanderson, G. C., 1966. The study of mammal movements: a review. Journal of WildlifeManagement. 30(1):215-235.

Sankar, K. 1994. Ecology of three large sympatric herbivore (chital, sambar, nilgai) with specialreference for reserve management in Saris tiger Reserve, Rajasthan. Ph.D. thesis, University ofRajasthan, Jaipur.

Sankhala, K. 1977. Tiger: the story of the Indian tiger. Simon and Schuster, New York.

Sankhala, K.S. 1978. Tiger, Rupa, Delhi.

Sankhala, K.S. 1993. Return of the tiger. Lustre Press, New Delhi.

Schaller, G. B. (1967). The deer and the tiger. Chicago: University of Chicago Press.

Schaller, G. B. 1972. The Serengeti lion. University of Chicago Press, Chicago.

Seidensticker , J. 1976. Ungulate populations in Chitawan Valley, Nepal. Biol. Conserv. 10: 183-209.

Seidensticker, J. & Mcdougal, C. (1993). Tiger predatory behaviour, ecology and conservation.Symp. Zool. Soc. Lond. 65:105-125.

Seidensticker, J. (1997). Saving the tiger. Wildl. Soc. Bull. 25: 6-17.

Seidensticker, J. 1976. On the ecological separation between tigers and leopards. Biotropica 8:225-234.

93

Seidensticker, J. 1986. Large Carnivores and the consequences of habitat insularization: Ecologyand conservation of tigers in Indonesia and Bangladesh. In Cats of the world: Biology,Conservation and Management, ed. S.D. Miller & D.D. Everett, pp. 1-41. Washington DC,National Wildlife Federation.

Seidensticker, J., Christie, S. & Jackson, P. (1999). Preface. In Riding the tiger: tiger conservationin human-dominated landscapes: xv-xix. Seidensticker, J., Christie, S. & Jackson, P. (Eds).Cambridge: Cambridge University Press.

Seidensticker,J.C. 1976. On the ecological separation between tigers and leopards. Biotropica8:225-234.

Simberloff, D.S. and L.G.Abele. 1976. Island biogeography theory and conservation practice.Science 191:285-286.

Smith, J. L. D. (1993). The role of dispersal in structuring the Chitwan tiger population.Behaviour 124: 165-195.

Smith, J.L.D. 1984. Dispersal, communication and conservation strategies for the tiger (Pantheratigris) in Royal Chitawan National Park, Nepal. Ph.D. thesis, University of Minnesota, St. Paul.

Soule, M.E. 1986. Conservation biology: the science of scarcity and diversity. 584pp

Springer, J.T. 1967. Some sources of bias and sampling error in radio -triangulation. J. Wild.Manage. 43: 926-935.

Sunquist, M E, 1981. The social organisation of tigers (Panthera tigris) in Royal ChitwanNational Park, Nepal. Smithsonian Contribution to Zoology. 336, 1-98.

Sunquist, M. E., Karanth, K. U. & Sunquist, F. (1999). Ecology, behaviour and resilience of thetiger and its conservation needs. In Riding the tiger: tiger conservation in human dominatedlandscapes: 5-18. Seidensticker, J., Christie, S. & Jackson, P. (Eds). Cambridge: CambridgeUniversity Press.

Sunquist, M.E. and F.C. Sunquist. 1989. Ecological constraints on predation by large felids. InCarnivore behaviour, ecology and evolution (ed.) J. L. Gittleman. 283-301.

Tamang, K.M. 1982. Population characteristics of the tiger and its prey. Ph.D. thesis. MichiganState University. East Lansing.

Terbough, J. 1976. Island biogeography and conservation: strategy and limitations. Science193:1029-1030.

Tester, J.R. and D.B. Siniff. 1965. Aspects of animal movement and range data obtained bytelemetry. Transactions of the North American Wildlife and Natural Resources Conference, 30.379-392.

Thapar, V. 1989. Tigers: the secret life. Elm Tree Books, London. 160pp.

94

Thapar, V. 1992. The Tiger’s destiny. London, Kyle Cathie.

Thomas, L., Buckland, S. T., Burnham, K. P., Anderson, D. R., Laake, J. L., Borchers, D. L. &Strindberg, S. 2002-a. Distance Sampling. In Encyclopaedia of Environ metrics. El-Shaarawi, A.H. & Peigorsch, W. W. (Eds). John Wiley & sons. Pp 1: 544-552.

Thomas, L., Laake, J.L., Strindberg, S., Marques, F.F.C., Buckland, S.T., Borchers, D.L.,Anderson, D.R., Burnham, K.P., Hedley, S.L., and Pollard, J.H. 2002-b. Distance 4.0. Release 2.Research Unit for Wildlife Population Assessment, University of St. Andrews, UK. http://www.ruwpa.st-and.ac.uk/distance/

Thompson, W. L., White, G. C. & Gowan, C. (1998). Monitoring vertebrate populations. NewYork: Academic Press.

Vanak, A.T.F. 1997. Movement Patterns of Radio-Tagged tigers in Panna National Park, MadhyaPradesh. M.Sc. Dissertation thesis, Saurashtra University, Rajkot, India. Pp. 47.Wemmer, C., J.L.D Smith and H.R.Mishra. 1987. Tigers in the wild: the biopolitical challenges.In Tigers of the world: biology, bio-politics, management and conservation of an endangeredspecies (eds.) R.L.Tilson and U.S.Seal. 396-405.

Wikramanayake, E. D., Dinerstein E., Robinson J. G., Karanth K.U., Rabinowitz A., Olson D.,Matthew T., Hedao P., Conner M., Hemley G. & Bolze D. (1999). Where can tigers live in thefuture? A framework for identifying high priority areas for the conservation of tigers in the wild.In Riding the tiger: tiger conservation in human-dominated landscapes: 255-272. Seidensticker,J., Christie, S. & Jackson, P. (Eds). Cambridge: Cambridge University Press.

Wikramanayake, E. D., Dinerstein, E., Robinson, J. G., Karanth, K. U., Rabinowitz, A. R., Olson,D., Mathew, T., Hedeo, P., Conner, M., Hemley, G. & Bolze, D. (1998). An ecology based methodfor defining priorities for large mammal conservation: the tiger as a case study. Conserv. Biol. 12:965-878.

Williams, B. K., Nichols, J. D. & Conroy M. J. (2002). Analysis and management of animalpopulations. San Diego: Academic Press.

Wilson, K. R., & Anderson, D. R. (1985). Evaluation of two density estimators of small mammalpopulation size. Journal of Mammalogy 66: 13-21.

Wright, B. 1996. The Tiger Poaching Crisis. CBI Bulletin. V4 (2): 23-28.Zar, Jerrold H., (1984). Biostatistical Analysis (Second Edition). Prentice-Hall, INC. EnglewoodCliffs, New Jersey 07632.

95

Appendix-I

Communities and Tiger Conservation:A holistic approach to development in and around

Panna Tiger Reserve

A Preliminary ReportJune 2005

ByVijay Kumar

Koustabh SharmaFaiyaz A. KhudsarR. S. Chundawat

Joanna Van Gruisen

BAAVAN: Bagh Aap Aur VAN

S-17 Panchsheel Apartment, A-1 Block Panchsheel Enclave,

New Delhi 110 017,email: [email protected]

Supported and Funded by:Global Tiger PatrolDiscovery Initiatives

96

COMMUNITIES and TIGER CONSERVATION : A HolisticApproach to Development In and Around Panna Tiger Reserve

Introduction Tigers are the largest of the living wild felids. They are at the top of the food pyramid and cansafely be depicted as the thermometers of any ecosystem. Since a healthy tiger population dependson a sound foundation underneath in the food pyramid, protection of tigers ensures health of theirprey and of the ecosystem, which in turn ensures water regime and the quality of floral richness ina particular area. A healthy and functional system is more beneficial to local communities thatsustain their livelihood on such forests than a poor quality and degraded forest habitat. Anonymous but systematic records have reported a drastic decline in the tiger population overthe last century. Poaching and habitat destruction are the two most significant factors responsiblefor this decline. In 1973, the then Prime Minister of India, Mrs. Indira Gandhi initiated an ambitiousproject to save the grandest of India’s natural heritage, the tiger. Project Tiger was initiated with anobjective of protecting tiger and its habitat and ensuring their long-term survival. Attempts were,and are still, made to provide protection to the tiger and its habitat through an extensive network ofprotected areas. This model had initial success but now is criticised for some of its serious fallouts:creating islands of protected areas without any corridors, alienating people from their ancestralresources and disputes over land use with the local inhabitants, these have created conflict withlocal communities. The protected areas have aimed at protecting our natural resources exclusively,in a way that involves little real interaction or bi-directional support to and from the people livingaround these areas. As a result of this exclusive approach, little has been gained long-term in termsof wildlife conservation. Rising conflicts with villages adjoining PAs is a cause for serious concernfor both wildlife conservationists and anthropologists working with the communities in these areas.The alienated villagers become potential allies to individuals, organisations and corporates withvested interests in the natural resources and in many illegal activities. Tigers are large territorial carnivores, which range widely and require large areas. Research hasrevealed that tigers have particularly large territories in Tropical Dry Forests such as in Panna NationalPark; they move great distances and their territories often extend beyond PA boundaries intounprotected tiger habitats dominated by a large human population. Lack of buffer areas, fragmentationand lack of corridors in forests result in an adverse situation for the tigers, as they are left with nospace for young individuals to disperse. At the same time, the territorial population of breedingindividuals face high risk when they foray out of the protected area in the routine course of theirterritorial movements. It is in these areas, outside the protected areas’ boundaries, where conflictexists, which makes this population highly vulnerable to existing threats. From past experience oftiger populations being lost from several of its strongholds, it is evident that the isolation model hasnot adequately achieved its objectives; this is true of the tiger, and also of other species. Habitat loss is a serious threat to tiger and associated biodiversity; it is critical to establishprotected areas as safe havens. Protected areas can have many management objectives beyondbiodiversity conservation, including community participation towards watershed development,sustainable livelihood, outdoor recreation and tourism, scientific research, and environmentaleducation and awareness. A Protected Area needs to be more than just the setting aside and guarding of a tract of land.Apart from management plans for protection of the fauna and flora, it is also essential that micro-plans are prepared for and with the communities residing at the periphery and inside the PAs,compatible with, and to ensure, protection of their biodiversity. Such plans need to address thevarious threats to the area and the biodiversity it supports, provide solutions for community needs

97

and for their appropriate and sustainable development. While environmentalists, biologists and wildlife conservationists play an important role indeveloping management/micro plans by addressing the ecological issues related to conservation,social scientists such as anthropologists, economists, and political analysts need to play an increasinglygreater role in helping protected area managers address the social issues related to conservation. Toensure protected areas’ success in conserving biodiversity, it is critical to understand the culturesand needs of the individuals and communities living inside and around the protected areas.Incorporation of communities as direct stakeholders in conservation can also extend conservationefforts beyond the borders of protected areas and has paid rich dividends in many parts of the world(Wells, 1996). Panna Tiger Reserve presently encompasses 543 km2; in the pipeline is another approximately200 km2 area of the Gangau Wildlife Sanctuary which is being added to the Tiger Reserve.Approximately forty seven villages are situated around the National Park and thirteen villages arestill inside the National Park’s boundaries. Villages inside the PA enjoy some privileges in terms ofgrazing, collection of minor forest products and restrained fuel wood collection. However, thesevillages are earmarked for relocation outside the PA. Some of the villages are already in the processof rehabilitation. The villages at the periphery of the PA appear to be the most underprivileged, mainly becausethey do not have access to the forest resources and development benefits have rarely reached them.Due to this, the standard of living in these villages is generally low and inevitably forest resourcesare used to fill the gaps; a growing population increases the damage caused by such activities on theadjoining forests. Repressive measures alone by the management, without alternatives provided,cannot lead to lasting solutions. Instead, they create conflict which continues to grow to unmanageableproportions. A holistic approach empowering the local communities is required for conserving theseforests while keeping in mind their economic, social and cultural development. Community-based natural resource management can empower communities through the effortsto manage available resources. Such management is based on the premise that those who are mostdependent on natural resources for their livelihoods have the greatest incentive to manage themresponsibly, and that they often possess an intimate knowledge of their local resources. Wherenatural resources are important for livelihood, empowering communities to manage these will haveenormous social, economic and environmental impacts. Involving communities in governance is anissue of direct relevance and can allow them to derive economic benefits from this improvedmanagement. It is important to generate alternate means of livelihood, reduce dependency of localcommunities on forests and make them active partners in conservation by channelling direct benefits. To develop a model where communities engage in supporting the management of local forests,wildlife, fisheries and water resources, a detailed understanding of the dynamics, demography andsocio-economic status of the communities is required, as well as some appreciation of thecommunities’ own views and perceptions of the existing problems. Baavan initiated this presentstudy in an effort to move towards such an understanding; the aim is to share the results of thissurvey, done with some participation of members of the local community, with the villagers so thata community conservation model can be developed with their active participation.

Area of interestThe 543 km2 of Panna Tiger Reserve in Madhya Pradesh is Tropical Dry Forest. It is situated

between coordinates 24°15'-24°20' N and 80°00'-80°15' E with an altitude ranging between 200mand 550m. The terrain of the area is typified by extensive plateaus and gorges. The unusual ‘step’topography gives the area three distinct levels: the Hinauta and Talgaon plateaus are the middle andupper levels respectively. The lowest ‘step’, the Ken river valley, splits the Reserve into two; steep

98

slopes, cliffs and deep gorges separate the different plateaus and the river valley. Panna receives an average annual rainfall of 1100 mm of which almost 60-70% falls betweenJuly and September. The area is lush green during and immediately after the monsoon; winters canbe characterized from October to January when temperatures may go down to as low as about –0.70C. A dry summer season follows until the next monsoon. Water is a major limiting factor duringthe summers when temperatures can rise to 47oC. The water table is peculiarly low and a largenumber of borings are either failures or have dried up within short durations.

A large cattle population of about 9,500 and over 6,000 humans live inside the Tiger Reserve.A much larger population of livestock and humans (37,500 cattle and 34,500 humans) surroundsthe Tiger Reserve, and depends largely on the forest for their daily requirement of fuel wood, grazingand on other minor forest produce. A monsoon, ‘rabi’, crop of wheat, gram or mustard is the mostcommon cropping pattern of the area. This forced single crop is due to the lack of water for irrigation.Waged labour, minor forest produce (MFP) collection and dairy are the other main sources of earninga livelihood in most of the villages.

Seven villages were selected on the basis of their geographic location with respect to PannaNational Park and proximity to the Hinauta village where the research base is situated. These villagesare typically constrained as they are surrounded by National Park and Sanctuary boundaries. Nospace to grow and few available resources put these villages in a high conflict zone: their needs aresufficed only by illegal wood and MFP collection, grazing, and mining.

Socio-Economic SurveyBefore initiating community-based conservation activities we considered it mandatory to collect

baseline socio-economic data on the target villages. Such preliminary surveys are essential forsuccessful and sustainable implementation of any developmental activity so that informed decisionsare made by the community. Once information from the survey is collated, analysed and simplified,we plan to translate it into Hindi and share it with the community to help them also to have a morecomprehensive understanding of the problems. The thorough socio-economic survey of all inhabitants of seven villages has consumed a largeamount of time but we hope it will help the communities to plan better. We aim for a largerparticipation over this planning process because we sincerely believe that such an approach willovercome some initial apprehension shown by the villagers during earlier meetings. The surveyobjective is to get a better insight into each village community’s composition and structure, theireconomic status, cattle strength, and dependency on the forests. Since the selected villages are justoutside the park, they do not enjoy the usual perks of land use and grazing as villagers inside thepark do. This has led to high amount of conflict because every activity which exploits naturalresources, such as grazing, fuel wood and MFP collection, is termed illegal. Their resources arelimited and legally they have no support for their high cattle populations, irrigation, fuel and othereconomic needs. The survey’s objective was to quantify activities that are based on natural resourcesand to gather information, which can help formulate appropriate action plans to reduce the conflict.

Information on basic demography parameters, employment, livestock population, cattle food,milk yield, firewood and its consumption, and minor forest produce was gathered from 602households of seven villages. The following section provides information on these villages, after asimple preliminary analyses of the data:

AnalysisPopulation Structure and Composition

The survey was design to collect information on the available workforce, requirements ofresources, community wise discrimination of available sources of employment.

99

The total population of these seven villagesis 3239 individuals in 602 families; the averagefamily comprises 5 to 6 members. Overallpopulation is close to unity with a slight biastowards males (1000 male: 953 female; this isbetter than the country average which is1000:933). Family size varies slightly amongdifferent communities. The largest families arefound in the Harijan communities, whereasAdivasis (Tribals) have the smallest families.The Muslim community showed a greatvariation: it had both smallest and largestfamilies (Figure 1). Very similar trends wereobserved among the different villages with very

little variation in family size (Figure 2).Adivasis form the largest community (52%),followed by OBCs (Other Backward Castes)(30%) and Harijans (11%). General and Muslimcommunities together account for 7% of thetotal population of these seven villages.Umrawan, Madaiyan and Bador villages havethe largest Adivasi population. Umrawan is anexclusively adivasi village. Bakchur comprisesonly of OBC community; and Kaimasan, whichis a very small settlement on the road, has amajority of Harijans who are otherwise in aminority throughout the other six villages.Hinauta and Bador, the two largest villages,

Average members

5.44 5.38

5.805.22 4.81 5.52

6.22

5.59

0.00

2.00

4.00

6.00

8.00

10.00

Bad

or

Bakc

hur

Dar

era

Hin

auta

Kai

mas

an

Mad

aiya

n

Um

raw

an

Tota

l

community

fam

ily s

ize

Figure 2. Average family size in the surveyed villages.

Average members

5.38

5.33

5.185.375.54

6.49

0.00

1.002.003.00

4.005.006.00

7.008.009.00

General Harijan OBC Tribals Muslim Total

community

fam

ily s

ize

Figure. 1. Average family size in different

communities in the surveyed seven villages

4% 2%10%5%

25% 11%

93%

28%

100%

22% 33%

7%

11%

64%

51% 44%

89%

100%

2%

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

Bador(119)

Bakchur(25)

Darera(86)

Hinauta(287)

Kaimasan(9)

Madaiyan(44)

Umrawan(32)

Muslim

Tribals

OBC

Harijan

General

Community wise population distribution

Figure 3. Village wise population distribution in the surveyed villages.

100

have a comparatively high mix of communities (Figure 3).

Community EconomicsLabour is the predominant employment

comprising of about 69% of the total workingclass. Hinauta, the largest village also has thelargest number of people employed inbusiness and labour. Contrary to a generalimpression, it is interesting to note from theresults of this survey that a very small numberof people depend on dairy for a living. When occupations are looked intocommunity wise, it is clear that the mainoccupation for Adivasis is labour and

agriculture. A small population of Adivasi community is dependent on cattle rearing and governmentjobs. OBC, which are mainly comprised of Yadav community, are largely dependent on livestockrearing and dairy related occupations and agriculture. But most of the milk production is for localconsumption. The rest of the population belonging to Harijan, Muslim and general caste are employedmainly in businesses and jobs, either forthe government or the diamond mine.

Settlements around forests havehistorically been dependent on thoseforests for their livelihood andsustenance. While setting aside theseforests as national parks and sanctuaries,the difficulties caused by instantly cuttingoff people’s dependence on them, needsto be recognised. Most people use fuelwood for cooking and for warming

Communitywise Occupation

0%

20%

40%

60%

80%

100%

Labour(n=476)

Agri (n=107) Cattle(n=45)

Business(n=39)

Job (n=24)

Occupation

prop

ortio

n

Muslim

Tribals

OBC

Harijan

General

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

Bador(132)

Bakchur(25)

Darera(103)

Hinauta(314)

Kaimasan(9)

Madaiyan(76)

Umrawan(32)

Village

Occ

upat

ion

Business

Job

Cattle/Milk

Agriculture

Labour

Figure 4. Village wise occupation in the surveyed villages.

Cattle number

0

100

200

300

400

500

600

General Harijan OBC Tribals Muslim

Cow

Ox

Buffalo

Goat

Pig

101

purpose despite the easy availability of cooking gasin some villages. Most households collect MFP andthis constitutes a significant chunk of their annualincome. Dependence on forests is extensive and theannual impact can be evaluated by the total biomassextracted annually.

Minor Forest Produce (MFP) is one of theprime sources of income for the population otherthan labour. Most of the households, up to 60%(361) of all the families surveyed (602), are involvedin MFP collection. Mahua, Madhuca indica (used

for brewing alcohol), Aonla/amla, Emblica officinalis (used for medicinal purposes and as a savoury)and Achar, Bucnania spp. (chirongi as dry fruits) are the main MFP collected from the Tiger Reserveand its neighbouring forests. The Adivasi community is mainly involved in MFP collection and tosome extent the OBC (Figure 5). Aonla and Mahua are the two most important MFP items collectedby them from the Tiger Reserve.

Achar, Mahua and Aonla are theprimary non timber minor forest products extractedfrom the forest. It is estimated that over 5.7 tons ofAchar is extracted from the forest during April andMay each year. Mahua flowers, which are extractedin the months of March and April, add up to amassive 36.5 tons. Aonla is the least expensive ofthe three MFPs but it is extracted in mammothquantity - 58.8 tons. The cumulative earning fromMFP collected in all seven villages is INR7,15,000.00 (About US$16,000.00). If considered per family income (consideringthat 60% of families are mainly involved in MFP collection), this comes to $45 (approximately INR2,000/- per family) which is a small amount; however the impact this collection causes in terms ofdamage and disturbance to the reserve is significant. It will be an important exercise to communicatethis information to the local community to understand the problem in a cost-benefit context. Theamount involved in MFP collection is not very large and if similar revenue can be generated throughalternate sources it may be possible to greatly reduce the biotic disturbance within the park. It willbe difficult to bring a complete stop to these activities but by providing alternatives and with aneducation awareness programme, its impact on the Tiger Reserve may be minimized.

Fuel wood, is the primary source of energy inthe surveyed villages. More than 597 (99%) of thehouseholds use wood. In Hinauta, despite easyavailability of cooking gas, only 44 families out of287 use this alternate energy source. The impact offuel wood demand on the forest is substantial asevery week 25 tons of wood is used by the villages.Although the impact of each village is proportionalto its population, it appears from the survey datathat the community structure does affect fuel woodconsumption. From the preliminary analysis it

Achar (kg/village)

Kaimasan (190kg)

3%

Madaiyan (487.5kg)

9%

Umrawan (785kg)

14%

Bador (917.5kg)

16%Bakchur (330kg)

6%

Darera (215kg)

4%

Hinauta (2790kg)

48%

Mahua (kg/village)

Hinauta (6030kg)

16%

Darera (3950kg)

11%

Bakchur (1390kg)

4%

Bador (14110kg)

39%

Umrawan (3690kg)

10%

Madaiyan (5820kg)

16%

Kaimasan (1580kg)

4%

Aonla (kg/village)

Kaimasan (2125kg)

4%

Madaiyan (6300kg)

11%

Umrawan (9000kg)

15% Bador (19262.5kg)

33%

Bakchur (750kg)

1%

Darera (0kg)0%

Hinauta (21375kg)

36%

102

appears that the Adivasi community is using morewood per capita than any other community and areclosely followed by harijans. This in turn affects theoverall impact of a particular village on the forests.This needs further analysis and exploration todetermine why there is higher consumption amongthese communities; then appropriate activities canbe planned to minimize the impact on the TigerReserve. The over all impact of a particular villageon the forest has two co-variates, viz. populationsize and community structure. Our approach tomitigate this problem has to be judiciously designed to address the issue both at a population andcommunity level. Further analysis is underway for assessment of these complex associations.

BIOGAS- an alternativeto Fuel-wood

Biogas is a clean andcheap fuel that contains55% to 70% of flammableMethane gas (CH4). It isproduced from cattle dung(gobar) through theprocess of decompositionin a specially designeddigester unit, which iscalled Gobar-Gas or Bio-

Gas plant. As a bi-product of this process, the productiveness, as manure, of the decomposed cowdung is increased up to approximately one and half times.

Benefits of Bio-gas plantApart from providing fuel and rich manure the introduction of Biogas plants to the local

community will release women and children from the pressure of (illegal) collection of fuel-woodfrom the Tiger Reserve and from preparing dung cakes. By removing smoke from the kitchen, itthus also reduces the risk of eye and lung ailments. Therefore, not only is it environmentally friendlybut a healthy and clean alternative to smoky fuel-wood. It also saves a substantial amount of cookingtime. If a Bio gas plant is connected with the sewage system of a house, it can improve the overallsanitation of the village. Besides cooking and illumination, the gas can also be used to run irrigationpumps.

Unit Type and CapacityDiscussions with families who have used or are using bio-gas plants have led us to conclude that thefloating model or KVIC model is the better one to use. Though this model costs more initially thanthe Dinbandhu model, which is more commonly promoted by the government, it is found to bemore successful in producing the gas on a regular basis. There are several problems associated withthe Dinbandhu model, one of which is the inconstant supply of gas. Because the chamber where thegas is collect is fixed and small, after 30 to 40minutes of use the pressure declines, affecting thesupply of gas to the burner. On occasions there is not even enough pressure to use the burner and theusers have to wait until the pressure builds up again. This causes interruptions in cooking and is not

Overall Firewood UseMadaiyan (1880kg)

7%Umrawan (2790kg)

11%

Kaimasan (355kg)

1%Hinauta

(8395kg)34%

Darera (4530kg)

18%

Bakchur (1205kg)

5%

Bador (5938kg)24%

12-1620-2510-121004

8-1214-187-9 753

5-810-134-6 502

3-44-62-3 251

Open grazingStall fed

Family sizeAnimals required per day Cow dung required (kg/day)

Production (m3/day)

12-1620-2510-121004

8-1214-187-9 753

5-810-134-6 502

3-44-62-3 251

Open grazingStall fed

Family sizeAnimals required per day Cow dung required (kg/day)

Production (m3/day)

103

appreciated by the user. Another, not uncommon, problem is the appearance of cracks in the chambercausing a leak; this is more likely to occur where the masonry work is of poor quality. On the otherhand in the floating model pressure remains constant throughout, until all the gas is utilized. Moreover,at times the plant can continue to produce gas for couple of days even when dung is not added orforgotten by the user. The floating chamber is either made of metal or more recently by fibreglass

material, which lasts for many years- on average 8-10 years - with verylittle maintenance. Its onlydisadvantage is the initial cost and alarger space requirement; even so wefind it to be the more successfulmodel. A comparison cost of differentsizes is provided in the table.

Networking with NGO, Individuals and Govt. OrganisationsIn the last seven months, several informal meetings were organized with local villagers with

an objective of bringing them together. This exercise was useful in constituting a committee inHinauta village. Several rounds of discussion took place between members of Baavan and the villagecommunities. The objective of conducting many such meetings was to develop a relationship oftrust between the two groups before plunging into concrete activities. While we succeeded inanswering some of the queries made by the villagers, it was also encouraging to find them volunteeringto take up responsibilities. Since any community development activity should have representationfrom all the communities within the village, we encouraged them to choose representative memberswithin themselves. The final group was formed through democratic interaction and discussions andwith proportional representation of different communities.

Visit to ChitrakootOne of the first activities undertaken under this project was in relation to cattle rearing and

milk production; after several discussions held with villagers, the village committee of Hinautanominated a group of representatives whom we took on a visit to Chitrakoot near Satna (in M.P.)where the Govansh Vikas Evom Anushandhan Kendra, Arogya Dham, Deendyal Research Instituteis situated.The following activities were observed and explained by the experts in details:1. Importance of Stall feeding

18003500n.a.n.a.5 or higher

18003500115154

180035001300096053

180035001150079602

18003500950058201

Generalcategory

SC\STFloating model (KVIC)

Dinbandhumodel

Subsidy amount

NABARD authorized unit cost Unit capacity (m3)

18003500n.a.n.a.5 or higher

18003500115154

180035001300096053

180035001150079602

18003500950058201

Generalcategory

SC\STFloating model (KVIC)

Dinbandhumodel

Subsidy amount

NABARD authorized unit cost Unit capacity (m3)

0.5-0.8%1%Potassium

0.5-0.8%1%Phosphorous

0.5-1%1.5-2.0%Nitrogen

Local fertilizerBio fertilizersNutritious constituent

0.5-0.8%1%Potassium

0.5-0.8%1%Phosphorous

0.5-1%1.5-2.0%Nitrogen

Local fertilizerBio fertilizersNutritious constituent

104

2. Indigenous cattle breed improvement3. Vermicompost4. Biogas as an ultimate source of energy5. Preparation of other compost using cow dung, urine and weeds (such as lantana, ipomoea andother herbaceous materials)

Villagers saw and discussed the merits of the ten different breeds of cows and bulls whichwere properly managed in the research centre. It was a day long visit which helped the villagers tounderstand the importance of bovine economy. It also helped in the selection of cattle breeds suitablefor the area. The idea was to explore improved and more productive animal husbandry practices andto introduce villagers to the concept of stall feeding of cattle for better dairy production with theobjective of reducing/removing the livestock pressure on the park. The visit also served to educateourselves and them on how local breeds can also produce more milk with very little investment;villagers were also introduced to the numerous means of utilising the by-products that can makethem economically secure with a reduced number of cattle. Discussions with scientists from theinstitution led to a better understanding of the benefits of stall-feeding and how an improvedmanagement of fewer cattle can enhance their gross income.

Similar meetings were also held in other target villages, where an introduction to theorganization and its objectives were explained to the villagers. The response of the villagers waswelcoming but at the same time they were a bit apprehensive about any such ventures. It wasevident from the discussions with them that this apprehension is a fallout of their bitter experienceswith similar previous initiatives that had eventually failed in absence of proper research and follow-up actions. It was also observed by us that part of their apprehension was due to the imposing natureof government policies; rather than based on community need and the realities of the field situation,most of the earlier initiatives had a top-down approach. Such impositions does not encourageparticipation and fails to generate required enthusiasm. Such imposed actions will eventually leadto failure of what might otherwise be a brilliant concept. The overall experience and feedback fromthese meetings with the villagers was encouraging, as more members of the villages wanted to visitChitrakoot to see for themselves the potential of improved production of dairy products from thelocal breeds and the techniques developed by the organisation there. In view of the enthusiasticreaction, we plan to take a few more such tours for other villages.

Visit to Tara GramTara Gram is an organizational setup situated near Orchha, in Tikamgarh district, part of a

well known national NGO, “Development Alternatives”. Tara Gram works at the community levelfor employment generation and empowerment of poor communities. They function as a hub withspokes of an efficient social network. The main objective of our visit was to collect information onbiogas. We also learned about various activities that are being conducted in the field of alternateenergy sources. Tara Gram has a range of activities aimed at generating employment and energywith an entrepreneurship approach. Different sections within Tara Gram comprise of charcoalproduction from waste wood, electricity generation from ipomoea (weed) and other biodegradablewaste, production of hand-made paper (recycled paper), biogas production for domestic use fromcow dung and other biodegradable waste, providing low cost illuminating and production of energyefficient cooking stoves (chulhas), preparing and marketing a range of confectionaries such as pickles,jams, marmalade, sauces and chips from locally available fruits and vegetables. It was found thatwhile biogas was successful in some villages, detailed discussions with their programme staffhighlighted the point that to make it a successful venture, consistent maintenance support for a fewyears is essential. The social network of Tara Gram is also involved in water conservation works,

105

which include designing and building check-dams to cater for water requirements and massplantations to serve a two-pronged purpose of raw material for various developmental activities.

The visit to Tara Gram opened up various possible avenues that were not known to us, andalso stimulated ideas of other possible activities, which may be undertaken. The visit and discussionswith their staff was an eye-opener and educational. With their vast experience in and around PannaTiger Reserve, their willingness to associate with our community based developmental project wasencouraging. We hope to incorporate their experience and expertise in initial building and maintenanceof the bio-gas (KVIC model) and improved Chulhas (fuel-wood stove).

Meeting other NGOs:Baavan has met and interacted with a number of other NGOs and organisations working

locally in related fields; Unnati and ‘Action for Food Production’ (AFPRO) Gwalior were two ofthese. Through them we were able make some very useful contacts, which can be tapped whenrequired in future. The district office of Madhya Pradesh Agro Industry Ltd in Panna also providedus access to technical information needed for building biogas plants, right from the installationstage to its maintenance and to efficient productive methods that are indispensable for its success inremote areas.

Information gathered about the support this organisation can provide in installation,maintenance, training and more importantly financial support to such an activity is crucial inexpanding our activities in these villages with their help. We are still in the process of contactingseveral other organisations working on dairy related activities. One such organisation is related tocheese production so that its possibility can be assessed in detail.

The Zila Panchayat Chief Executive Officer’s (CEO) response to our initiatives was verywarm and he showed keen interest in our objectives and promised his support in making this projecta reality. He even assured financial support in the installation of biogas plants in the villages. Hissupport shall be crucial in assuring the holistic development of the villagers. The newly electedPresident of Zila Panchayat was also very keen to give a helping hand and assured a full cooperationwith the organization in its pursuit.

understanding tiger ecology in the tropical dry forest of

Documents