Vipera Ursinii Rakosiensis
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Lifehistory ,population characteristicsand conservation of the Hungarian meadow viper ( Vipera ursiniirakosiensis ) Beáta Ú jvári 1,Zoltán Korsós 1 ,Tamás Péchy 2 1 Department ofZoology, Hungarian Natural HistoryMuseum, Baross u.13,H-1088 Budapest, Hungary e-mail: [email protected] 2 HungarianOrnithological and Nature ConservationSociety, Kö lt" ou.21,H-1121 Budapest, Hungary Abstract. Duringa continuouspopulation study of Viperaursinii rakosiensis ,79specimens were measured in the eld,between 1993and 1997. Body length and body mass ofsix juveniles were comparedto those of 43 more specimens whichwere keptduring their rst winterin terraria. Signicant differences were foundin favour ofthe juveniles kept in captivity, whereas acomparisonwith earlier data from1953 showed a strongdecline injuvenile body mass. Based onbody size, three age classes couldbe distinguished in the study population. Theannual activity and reproductive cycle ofthe Hungarian meadow viperis describedfor the rst time. A comparisonto other European populations of Orsini ¢sviperrevealed aremarkablyhigher clutch size in Vipera ursiniirakosiensis ,whichis explainedby the larger bodysize offemales inthe particular population. Introduction TheHungarian meadow viper ( Viperaursinii rakosiensis Méhely, 1893) is a small- sized( totallength is about 60 cm) lowlandsteppe form ofthe Viperaursinii species group.Since the Viperaursinii speciescomplex is explained in different ways by the modernmorphological and biochemical research results, the systematic status of Vipera ursiniirakosiensis isstillunresolved, and the subspeci c rankcan only be consideredas provisional(V anceaet al., 1985; Joger at al., 1992; Nilson et al., 1993). TheHungarian meadow viper ( Viperaursinii rakosiensis )isone of themost threatened snakesin Europe(Taká cs et al.,1987; Corbett, 1989; Korsó s, 1992). The former distribu- tionof themeadow viper included the easternmost part of Austria, Hungary, Transylvania (Romania),and northern Bulgaria (Dely and Janisch, 1959). Populations today only exist intworegions of Hungary( inthe Great Hungarian Plain between the rivers Danube and Tisza,and in the Hansá g NatureReserve, northwestern corner of the country), from all theother territories it is mostprobably extinct (Korsó s, 1992; Pé chy et al., 1996). Since c KoninklijkeBrill NV ,Leiden,2000 Amphibia-Reptilia21: 267-278 ® 268 Beáta Újvári, Zoltá n Korsós, Tamá s Péchy 1974,the meadow viper has received full legal protection in Hungary where and since thenit has been one of the most strictly protected species. Theoretical nature conserva- tionvalue of onespecimen is 500,000 HUF (ca.1,960 EUR). Parts ofthedistribution area arealso protected; however, most of thisarea is notgovernmental property, which poses greatproblems for conservationto be effectivethrough habitat protection. The animal is listedin the 1996 IUCN RedList of ThreatenedAnimals as “ threatened”, appearson the CITES AppendixI andthe Berne Convention Appendix II. Eventwo recommendations oftheCouncil of Europeemphasise its strict protection in Hungary.It isalso included in theprogrammeof theHungarian National Biodiversity Monitoring System ( Korsós, 1997). Thesensitivity of thissnake to habitatalteration and human disturbance is quite dramatic. Populationsare subject to short term ( coldwinter, high soil water level, etc.) as well as longterm natural threats (isolation, genetic drift, inbreeding, etc.). However, the meadow viperis immediatelyand most signi cantly threatened by humanagricultural activities, like intensivegrazing, burning, machine mowing, etc ( seemore details in Korsó s andFü lö p, 1994;Pé chy et al., 1996). The biologyof theHungarianmeadow viper is poorlyknown, in partdue to itsrarity in nature. Inthe frame oftheconservation program of the Hungarian meadow viper a continuous populationstudy in theKiskunsá g area( southeastof Budapest,Hungary) is beingcarried outsince1993. The goalof thisproject was tocollectas many data as possibleabout the life historyand population characteristics of thisthreatened snake. The program also included apopulationreinforcement experiment with the support of the Hungarian Ministry of Environment( Nechayand Pé chy, 1994), which aimed at the improvement of thesurvival chancesof aselectedpopulation by helpingthe juvenile specimens to get over their rst winterin captivity. Theaim of thispaper is tosummariseresults on theannual activity of thespecies, with acomparisonof data obtained from captivityand from thewild. The age structure, and annualactivity cycle is describedof the selected population, and compared to two other EuropeanAlpine populations of Viperaursinii . Materialand methods Altogether,79 meadow vipers (25 males, 48 females, 6 juveniles)were measuredin the Dabas-Gyón studyarea between 1993 and 1997. The typical vegetation of the habitat consistedof awet,closed grassland ( dryingwet meadow = Molinietum)plantcommunity ofuneven structure, characterised by the grass species Moliniacoerulea , Schoenus nigricans, Chrysopogongryllus and Stipa sp.The structure of thevegetation is arranged intomicrolayers, interspaced by tussocksof grassof differentages. The densityof possible preyitems (insects: large-sized orthopterans, and young lizards: Lacertaagilis , L. viridis and Podarcistaurica )isoften high. For moredetails see Ú jvári andKorsó s (1997). Life historyof Viperaursinii rakosiensis 269 Table 1. Numberof captures of V.u.rakosiensis inDabas-Gyó n. Males Females Juveniles Recaptures 1993 7 5 27 (7)a 1 1994 7 14 14 (1)b 2 1995 11 22 2c 4 1996 – 5 4 1 1997 – 2 2 1 Total 25 48 49 (8) 9 a Bornfrom three females incaptivity; vedied immediately, two during the winter period. Twenty-seven were released inMay1994. b Elevenwere caughtin September 1994, four were bornfrom a female incaptivity. One juvenilefemale died duringthe winter. Fourteen juveniles were released onthe original spot in May 1995. c Caughtin August 1995 and released inMay 1996. Dataalso originated from apopulationreinforcement program. In autumn 1993 and 1994,gravid females were collectedand kept in terraria until they gave birth. The young were housedin an enclosed part of theZoological Garden, Budapest until the subsequent spring.The furnishingof theterraria imitated natural habitat ( grassand soil were collected ontheoriginal locality), and to minimize the disturbance of theanimals only one person was allowedinto the room during feeding times. To avoid the association of food with humanpresence no record was keptof the exact food consumption of juveniles. Insects (large-sizedorthopterans) and newborn rodents were providedad libitum, i.e. for all neonatesmore food was availablethan they could utilize. These 30 juveniles,together with 13morecaught in theautumnand handled in thesame way during winter, were releasedin goodcondition in Mayof thesubsequent year at the original collecting locality (table 1). Bodyweight and total length at the juveniles maintained in captivity were measured oncein everymonth during the winter.They were alsomeasured at thedate of theirrelease, andthe data were comparedto those by Lányi ( 1957).In the course of ourstudy not all morphologicaldata were consequentlynoted, hence the number of specimensincluded in ananalysis of agivenmorphological character might differ. Results Bodysize comparison Changeof body mass andbody length of the meadow viper juveniles during their rst winteris plotted in gures1 and2 (repeatedmeasures of the same specimens on the sameday). For averagebody mass andlength, there were nostatisticaldifferences between neonatefemales ( x¯ = 3.0 g, s = 0.13, x¯ = 150.3 mm, s = 1.56)andmales ( x¯ = 2.9 g, s = 0.35, x¯ = 148.0 mm, s = 2.5).Using regression analysis ( Sutherland,1996) thegrowth rate of the captive specimens showed an allometric relation: for bodymass thecorrelation was exponential( x = month, y = 2.32e0.16x, r 2 = 0.9665, t = 15.19; 270 Beáta Újvári, Zoltá n Korsós, Tamá s Péchy Figure 1. Bodymass ofjuveniles during winter ( opencircles = specimens keptin terraria; lleddots = measured inthe eld,after hibernation). Table 2. Comparisonof captiveand in situ specimens after their rst winter( mean ± standarddeviation, sample size). Bodymass [g] Bodylength [mm] Keptin terraria 15.7 ± 2.97 (18) 287.0 ± 27.88 (18) Measuredin the eld 6.8 ± 1.91 (9) 203.8 ± 28.39 (10) P < 0.1%, df = 8),whereas for bodylength was linear( x = month, y = 13.16x+ 127.39, r 2 = 0.9701, t = 16.11, P < 0.1%, df = 8).Measurements of thecaptive specimens at thedate of thereleasewere comparedwith those measured in theeldin May ( table2; g. 1and2: opencircles and lleddots, respectively). The differences signi cant ( bodymass: t = 7.41, P < 0.1%, df = 23;body length: t = 6.64; P < 0.1%, df = 23).The terrarium specimensgained an average8.9 g and84 mm increaseduring the winter. Acomparisonof ourmeasurements on juvenilesborn in with literature data from 1953 (Lányi, 1957) show a conspicuousdifference in juvenilebody size. While the body mass ofthejuveniles forty years ago appeared to be signicantly higher than today ( t = 3.07; P < 1%; df = 23; g.3),their total length was foundto be smallerthough on a lower signicance level ( t = 2.17; P < 5%; df = 25; g. 3). Age classes Onthe basis of the body mass-body length ratio as well as the capture-recapture data, threeage groups can clearly be identied in theDabas-Gyó n population( g. 4).The rst Life historyof Viperaursinii rakosiensis 271 Figure 2. Bodylength of juveniles during winter ( opencircles = specimens keptin terraria; lleddots = measured inthe eld,after