JOURNAL OF THE BOMBAY NATURAL HISTORY SOCIETY August 2001 Vol.98 No. 2 ESTIMATING THE DENSITY OF PORCUPINES IN SEMI-ARID SARISKAVALLEY, WESTERNINDIA' DiwakarSharma2 (With three text-figures) Key words: Indirect estimates, direct estimates, dung decay, pellet groups, defecation The population of the Indian crested porcupine (Hystrix indica) was estimated over a periodof3months(May-July 1989)insemi-aridSariskavalleyintheSariskaTigerReserve, Rajasthan. The methods included count ofpellet groups in four transects each in three vegetation types, and direct counts ofanimals in a vehicle at night. The data collection using these two approaches was repeated 17 times. Calculation of density for indirect estimatesinvolveddataonthepelletdecayrateandthedefecationrate(ofcaptiveanimals). The animal density was estimated to be 8.8 ±2.4 animals/ sq. km through direct count, whilepellet group count provided an estimate of12.4 animals/ sq. km with a variation of 0.9 to 24.9 animals/ sq. km in different habitats. The influence ofsome factors such as possibleuseoflatrinesitesandslowdecompositionrateondensityestimatesarediscussed. Introduction feeds on crops (Alkon 1983, Gutterman 1987, 1988) and on the bark of trees (Choudhry and To estimate the population density of Ahmad 1975, SharmaandPrasad 1992). Inspite animals in the wild, quantifying faecal matter ofits economic importance, few estimates ofits abundance, decay and defecation rate is population density in the wild are available. considered to be useful and convenient (Neff Beingnocturnal,smallinsizeandshyofhumans, 1968, Rowland etal. 1984). Although anumber itishardtoobserveandverydifficulttoestimate of studies have been conducted elsewhere its population. Therefore, methods involving (Putman 1984, Koster and Hart 1988), in India indirect evidence can be very useful for onlyafewstudieshaveestimatedthepopulation estimation ofitspopulation. Duringthe summer of wild animals using this method (Sale et al. of 1988, the last of three consecutive drought 1990, Dekker etal. 1991). years,comparativelylargenumbersofporcupines The Indian crested porcupine Hystrix were seen in Sariska valley in the Sariska Tiger indica Kerris a nocturnal animal thatreportedly Reserve. This was a unique opportunity to estimate itspopulationusing direct sighting and 'AcceptedJanuary,2001 faecal abundance. 2WildlifeInstituteofIndia,P.B.#18,Chandrabani The aim of this study was to estimate DehraDun248001,Uttaranchal,India. porcupinedensityusingpelletgroupdensity, and PresentAddress:GujaratEcologicalSociety, 5,GoldenApartment,Subhanpura, to compare the results with that obtained using Vadodara390023,Gujarat,India. direct observations. JOURNAL BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG.2001 161 ESTIMATINGTHEDENSITYOFPORCUPINES Study Area temperature up to 47 °C) and monsoon (July- October). Average annual rainfall is c. 650 mm, Sariska Tiger Reserve (27° 20' N and 76° mostofwhich fallsbetweenJulyandSeptember. 25' E) is a dry deciduous forest (Champion and ThestudywasconductedbetweenMayand Seth 1968) located in the semi-arid tract of July 1989 in the Sariska valley (Fig. 1). The Rajasthan in western India. The weather is dry vegetationofthestudysitesconsistedofZiziphus and hot during the day throughout the year, scrubland, mixed woodland and Ziziphus barring the monsoon when it is hot and humid. woodland (Sharnia and Prasad 1992). The hills There are three distinct seasons: winter surroundingthe valleyhadforests dominatedby (November-February, with night temperature as Anogeissuspendula and were little used by the low as 3 °C), summer (March-June, day porcupine(SharmaandPrasad 1992). Therewas Fig. 1: Sariska valley showing the study sites 162 JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG. 2001 ESTIMATINGTHEDENSITYOFPORCUPINES very little ground cover and litter in the three Where, vegetation types during the study period. Y = mean pellet group density (no./sq. km) The major mammals in the Sariska r= mean decay rate per day National Park include tiger Panthera tigris D = mean defecation rate per day ( ), leopard P pardus), nilgai Boselaphus Pelletgroupdensity: Porcupinesdefecate ( . ( tragocamelus), sambar(Cei~vus unicolor), chital spindle shaped pellets, which are about (Axisaxis),wildpig(Susscrofa)andfour-homed 1-1.5 cm in diameter and 3-5 cm in length. The antelope (Tetracerus quadricornis). The density pellet number may range from 5-25 in a pellet of animals, particularly of ungulates, is higher group. Pellets groups were counted in strip in Sariska valley, due to better habitat, than in transects in the three vegetation types, selected other areas ofthe Tiger Reserve. under stratified random sampling. Four strip m transects 500 x 2 wereplacedparallelto each m Methods other at 100 intervals in each vegetationtype (Fig. 1). Each transect was visited 17 times Direct Density Estimates during May-June. The number ofpellet groups Since porcupines are nocturnal, they were thatwereavailableattheendofthestudyperiod countedatnightusingvehicletransects(Rodgers after their accumulation and decay in each 1991), along a 10 km long road (Sariska to vegetation type was used to calculate the pellet Kalighati), in Sariska valley. In June and July, group density i.e. the number ofpellet groups the census was done 17 times, between 2000 to per sq. km. 2030hrs,by fourobservers, inajeep drivenata Pellet decay rate: Investigations of the uniformspeed of20km/hr. The animals seen in decay rate ofpellets were carried out from the theheadlights ofthe vehicle were recorded. The first week ofMay 1989 to the second week of m fieldcoveredbytheheadlightswas 10 oneither July 1989. Fresh pellet groups were located side of the road. Porcupine density (D) was between May 10 and June 30. Eight groups of calculatedusingthefixed-widthtransectmethod freshpellets encountered were marked, counted (Rodgers 1991). andtheirconditionmonitoredonceaweek. Two N periods i.e. summer (May to mid-June) and D = pre-monsoon(mid-Junetomid-July)werechosen 2.L.B to compare the seasonal decay rates. Where, The following equation was used (Barnes N =number ofanimals observed, and Jensen 1987, Sukumar et al. 1991) to = L length ofthe transect in km, and estimate the decay rate for pellet. B = width on one side ofthe transect in km. ln(N)-ln(N = t tl) Indirect Density Estimates r Porcupine density (N) based on pellet t group count, defecation and decay rate was Where, calculatedusing the following equation (Barnes r= mean decay rate ofpellet groups per day, and Jensen 1987) N =number ofpelletgroups ofone week, Y.r N = the number ofpellet groups ofprevious N = wee]k, and D t= time (in days). JOURNAL BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG.2001 163 , ESTIMATINGTHEDENSITYOFPORCUPINES Defecation Rate: Two porcupines were Table2 observed in the National Zoological Park, New SUPMOMRCEURP(IMNAEYPTELOLMEITDD-JEUCNAEY)AINND Delhi for 15 days, one in Sakkarbagh Zoo, PRE-MONSOON(MID-JUNETOMID-JULY) Junagadh, for 19 days (in 1992) and another in SayajibaugZooinVadodarafor21 days(in2000) S.No. Time(days) Condition Summer Pre-monsoon and their pellet groups counted. The defecation ratewascalculatedasthenumberofpelletgroups 1 < 1 < 1 Fresh,moist,coveredwith defecatedperday. mucous,intact. Data was analysed using computer 2 1-7 1-7 Intact,butdry. 3 8-30 8-15 Drywithcracksover software. Non-parametric statistical tests were thesurface;morethan75% conducted for significance of results. pelletsdistinguishable. Kruskal-Wallis tests were used to determine 4 30-45 15-30 Crackswidenandcolour significance of variation among the three becomesdark;about50% ormorestilldistinguishable. vegetation types, while Mann-Whitney U 5a 30-45 Allpelletsdisintegrate. statisticswereusedtodeterminethesignificance 5b >45 Lessthan50% ofvariationbetween the two seasons. distinguishable. Table1 decay rates of different pellet groups within a DIRECTANDINDIRECTDENSITYESTIMA1uSFOR season. Trampling by large mammals such as PORCUPINEINDIFFERENTHABITATS wild pig, sambar, chital and nilgai accelerated Vegetationtype Pellet Animal Shrub the process. On the other hand the pellets not density density density (No./km2 trampled by large mammals were present even ) (No./km2) Direct Indirect (No./ha) e6n0vidraoynsmenatf.teNrobteeirnmgitedaecfteicoantweadsionbsaermvoedisitn Ziziphus scrubland 9975 2.35±2.16 24.9 411 ±114 summer. Ziziphus The pellet decay rate is shown in Figs 2 woodland 350 6.56±1.04 0.9 137±63 and 3. Table 3 shows the decay rate for various Mixedwoodland 5075 17.6±4.02 12.7 79±55 Average 5133 8.8±2.4 12.8 206±77 pellet groups monitored during summer and pre-monsoon. The average decay rate during summer was 0.005 per day. This rate increased Results to more than twice, 0.011 per day during Pellet Group Density pre-monsoon. The difference in the mean decay There was a significant difference in the rates between summer and pre-monsoon was pellet group density among the three vegetation significant(M-WtestsZ=2.31, 2 tailedp<0.02, types (K-W tests x2=16.15, p<0.001, n=34) n = 27,19). For calculation ofanimal density, (Table 1). on,ly the decay rate during summer (May-June) are considered, as pellet group densities were Pellet Decay Rate estimated only in this season. The pellet condition was categorised into five agegroups (Table 2). Pellets observedfrom Defecation Rate June onwards (afterpre-monsoon showers) also Inthewild,thepelletswerespindleshaped disintegrated through the same categories, but and discrete; while in the New Delhi zoo the the decay was faster in the second category. pellets in a group were not as discrete. In Results indicated a large variation in the SakkarbaghZoo(Junagadh)andSayajibaugZoo 164 JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG. 2001 ESTIMATINGTHEDENSITYOFPORCUPINES Days —— PelletGroup 1 PelletGroup 2 PelletGroup 3 Fig. 2: Decay rate ofporcupine pelletsduring summer Fig. 3: Decay rateofporcupine pelletsduring pre-monsoon JOURNAL BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG.2001 165 ESTIMATINGTHEDENSITYOFPORCUPINES DECAYRATESOFVTAaRblIeO3USPELLETGROUPS roadwas similartothatawayfromit. Therefore, MONITOREDDURINGSUMMERAND it was presumed that porcupine movement and PRE-MONSOONPERIODS distribution along the roads would be similar to Pellet Summer Pre-monsoon those away from the road. The small width (20 (Maytomid-June) (mid-Junetomid-July) m) of the transect precluded the chances of GroupNo. Pellet Decay Pellet Decay missing animals on it. Within each vegetation group rate group rate type, density estimates from vehicle census at size (perday) size (perday) night had low variation, except in the case of 1 17 0.005 5 0.010 Ziziphus scrubland. Further, the overall low 2 8 0.004 23 0.012 3 11 0.011 11 0.014 variation in average density of the animals in 4 10 0.020 the study area was considered good to compare 5 11 0.007 Mean 0.005 0.011 indirect estimates with it. The higher decay rate in pre-monsoon (Vadodara), because of near natural diet, the season (mid-June to July) was due to the action droppings were similar to those in the wild and ofdungbeetles andtermites. This startedwithin were included for calculations. The diet a week afterthe first shower inthebeginning of (includingaccesstowater)incaptivityisdifferent June. However, on hard (red soil) and bare from that in the wild, hence the defecation rates groundthedisintegrationbytermitestooklonger should be taken only as an approximation. If time. possible, the defecationrate shouldbeestimated Freshpellets were not monitored after the innaturalorsemi-naturalconditions. Defecation onset of monsoon in July because of rapid rate ofporcupine in captivity varied from once disintegration. Often within a day or two about to thrice a day. The average defecation rate was 50% or more of the pellets changed into an 2 ±0.15 per day (n = 40). amorphousmass.Further,denseundergrov±hdid notpermit an easy search offresh pellets. Since Ai^imal Density theweatherisdryfor8months,thedatacollected The results showed a difference between duringthisperiodisconsideredtobemoreuseful. the direct and indirect estimates of density in The observed differences in density each vegetation type (Table 1). The overall estimates for the two methods were perhaps density estimated by direct observation was 8.8 relatedto the shrub coverinthe three vegetation ±2.4 animalspersq.km,whiletheestimateusing types. Earlier results (based on debarking, indirect method was 12.9 animals per sq. km. Sharma and Prasad 1992), show that the porcupinespreferred mixedwoodland,Ziziphus Discussion woodland followed by Ziziphus scrubland as habitats. Onewould,therefore, expecttheanimal The road used for vehicle census at night abundance and pellet density to follow this passed through three different vegetation types, pattern. While the direct counts conformed to each of which differed from the others in thispattern,thepelletdensityandindirectcounts structure, composition, density andutility to the followed the reverse trend. This suggests that porcupine (Sharma and Prasad 1992). This while the porcupines preferred to feed in mixed resultedinvariationinporcupinedensityamong woodland and Ziziphus woodland, they spent thevegetationtypes.Duetotheimpactofdrought substantial time in the dense undergrowth of and livestock grazing, the vegetation along the Ziziphus scrubland. In Ziziphus scrubland, the 166 JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG. 2001 1 ESTIMATINGTHEDENSITYOFPORCUPINES shrub density was the highest, followed by only, while in captivitythe animals defecatedin Ziziphus woodland and mixed woodland (Table the corners of the cages. This indicates the 1). Dense shrub provides escape cover for the possibilityofuseoflatrinesitesbytheseanimals. porcupine (Prater 1993). Ifso, can this method be used for other animals This pilot study attempted to validate the that use latrine sites? methods used for indirect animal density Moreover, the semi-arid climate seems to estimates. Eberhadt andEtten (1956), Etten and have played an important role in substantially Bennet (1965), Neff(1968) and Putman (1984) slowingdownthedecayrateofpellets. Pelletsof have comprehensively reviewed the limitations chitalandsambardefecated inthepreviousyear andsourcesoferrorinsucha study. Asrevealed have been observed in Gir Wildlife Sanctuary in this case, the differences in density estimates and National Park in Gujarat by this author, between direct and indirect evidence strongly indicating their slow decay rate in similar emphasise the need for aproper stratification of conditions. These aspects require more research habitat,preferablyafterreconnaissance,followed and,perhaps, the inclusionofa correctionfactor by adequate sampling (Grieb 1958). The results inthecalculationofdensitiesinsuchconditions. indicate that to estimate the population of nocturnalanimalssuchasporcupine, oneshould ACKNOWLEDGEMENTS study all habitats including those that may be avoided by the animal (Neu etal. 1974). I thank the Chief Wildlife Warden, Even though most of these factors were Rajasthan, forpermissionto work inthe Sariska takenintoaccount(Sharma 1989)thedefecation Tiger Reserve and Mr. H.S. Panwar, former behaviour of porcupine may have played an Director, Wildlife Institute of India for logistic important role in the indirect density estimates. andfinancial support. Dr. AjithKumar, Dr. S.P. The droppings were clumped, andundernatural Goyal,M.V.KattiandT.R. ShankarRamangave conditions were observed at certain locations critical suggestions on the manuscript. ReferENCES Alkon,P.U.(1983): ForagingecologyofIndianCrested ofIUCN/SSCAsianElephantSpecialistGroup,pp. Porcupine{Hystrixindica)inNegevhabitats.Final 54-64. Report.IsraelAcademyofScience. 14pp. Eberhardt, L. & R.C. Van Etten(1956): Evaluation of Barnes, R.F.W. & K.L. Jensen (1987): How to count thepelletgroupcountsforcensusingdeer.J. Wildl. elephants in forest. IUCNAfrican Elephant and Manage. 20:70-74. RhinoSpecialistGroupTechnicalBulletin I: 1-6. Etten,R.C.Van&C.L. Bennet(1965):Somesourcesof Champion,H.G.&S.K.Seth(1968):Arevisedsurveyof errorinusingpelletgroupcountsforcensusingdeer. the forest types of India. The Manager of J. Wildl. Manage. 29:723-729. Publications,Govt,ofIndia,Delhi,404pp. Grieb,J.R.(1958):Wildlifestatistics.ColoradoGame& Choudhry,M.I.&A.Ahmad(1975):Trialsofpoisonous FishDepartment.96pp. gases and baits against porcupines. Pale J. For. Gutterman, Y. (1987): Dynamicsofporcupine(Hystrix 25(2):46-50. indica) digging: Their role in the survival and Dekker, A.J.F.M., S. Dawson & A.A. Desai (1991): An renewalofgeophytesandhemicryptophytesinthe indirectmethod forcounting Asian Elephants in Negev Desert Highland. Israel J. Botany 36: forests. In: Censusing Elephants in Forest. 133-143. Proceedings ofan International Workshop. (Eds Gutterman, Y. (1988): An ecological assessment of Ramaknshnan, U.,J.A. Santosh,&R. Sukumar). porcupine activity in a desert biome. In: Southern India, 2-10 January, 1991. Technical Ecophysiologyofdesertvertebrates. Eds Ghosh, & ReportNo.2.AsianElephantConservationCentre P.K. I.Prakash,ScientificPublications,Jodhpur. JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG.200 167 . ESTIMATINGTHEDENSITYOFPORCUPINES Pp.289-363. Sale, J.B., A.J.T. Johnsingh & S. Dawson (1990): Koster,S.H.&J.A.Hart(1988): Methodsofestimating Preliminary Trials with an Indirect Method of ungulatepopulationsintropicalforests.Afr.J.Ecol. Estimating Asian Elephant Numbers. Report for 26: 117-126. IUCN/SSCAsianElephantSpecialistGroup. Neff, D.S. (1968): Thepelletgroupcounttechnique for Sharma, D. (1989): Spatial and temporal patterns in biggametrend,censusanddistribution:Areview. debarking by Indian Crested Porcupine Hystrix ( J. Wildl.Manage. 32:597-614. indicaKerr)in SariskaNational Park, Rajasthan. Neu,C.W.,C.R.Byers&J.M.Peck(1974):Atechnique M.Sc. Dissertation, Wildlife Institute of India, foranalysisofutilizationavailabilitydata.J. Wildl. Dehradun,88p. Manage. 38:541-545. Sharma, D. & S.N. Prasad(1992): Tree debarking and Prater,S.H.(1993):TheBookofIndianAnimals.Fourth habitatusebyporcupine(Hystrixindica Kerr) in impression. Bombay Natural History Society, SariskaNationalParkinWesternIndia.Mammalia. Bombay,pp.215-217. 56(3):351-361 Putman,R.J.(1984):Factsfromfaeces.MammalReview Sukumar, R., K.S. Verman, J.A. Santosh & U. 14(1):79-97. Ramakrishnan (1991): Methods of estimating Rodgers,W.—A.(1991):TechniquesforWildlifeCensusin elephant numbers: A review. In: Censusing India AfieldManual.WildlifeInstituteofIndia Elephants in Forests. Proceedings of an Dehradun. 82p. International Workshop (Eds Ramakrishnan, U., Rowland,M.M.,G.C.White&E.M.Karlen(1984):Use J.A. Santosh, & R. Sukumar). Southern India, 2- ofpelletgroupplotstomeasuretrendsinestimates 10January, 1991,Technical ReportNo. 2. Asian in deerand elk populations. Wild. Soc. Bull. 12: ElephantConservationCentreofIUCN/SSCAsian 147-155. ElephantSpecialistGroup,pp2-29. 168 JOURNAL, BOMBAYNATURAL HISTORYSOCIETY, 98(2), AUG. 2001