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HJvrix,(n.s.) 10 (1) (1998): 17-30

RE-INTRODUCTION AND PRESENT STATUS OF THE LYNX (LYNX LYNX) IN

URS BREITENMOSER, CHRISTINEBREITENMOSER -WURSTEN AND SIMONCAPT

SwYS Lyzs Project, ~Tllettengiissli4, CH-3074 Muri, Switzerland

ABSTRACT - A lynx recovery programme started in Switzerland in 1970. From 1970-76. at least 14 lynx were translocated from the Carpathian into the Swiss . Another re-introduction took place in the , but no corridors exist as a connection between these two popula- tions in Switzerland. The devclopmcnt of the populations was not monitored at first. In 1980 sys- tematic research was initiated. which gradually cvolved into the'SM,iss Lynx Projwl. Not all releases were successful, but the re-introduction in the northcm and founded a population that covered an area of some 4000 kin' in 1981. In the wcstem Swiss Alps. lynx moved into Italian and French territory. Towards the eastern Swiss Alps. the expansion was slower and ceased about ten years ago. During the last five years. there even has been a reduction of the arca occupied. Today, the pop- ulation covers an area ol about 10000 kin2 in the Swiss Alps, of which 50% is suitablc lynx habitat. Based on sizc and overlap of average home ranges of radio-tagged lynx. the population was cstimat- ed to include soinc SO adult residents. At present, the growth rate of the population appears to bc too low to allow a further expansion in range. It is uncertain whether recruitment is sufficient to com- pensate for the high losses among resident adults induced by traffic accidents and illegal killing Kq-Mwds: Lym lyn.~,re -introduction, status. monitoring, depredation, mortality, Switzerland, Alps

INTRODUCTION west (Hell 196l)-and rumours about lynx migrating through the Alps (Schrnidt 1960) The lynx population of Switzerland was ex- directed wider attention to this large felinc. terminated during the 18th and 19th cen- In 1962. lynx and brown bear were given le- turies. This was a period when Switzerland gal protection in Switzerland. As a result, faced ecological disaster: Many of the plans of re-introducing the lynx were seri- wcrc destroyed or in an alarming ously discussed. and studies on the decline state, the wild ungulate species were eradi- of the species in Switzerland creatcd further cated (CPI.I,USelaplius. Caprvolrrs mapeo- support (Schauenberg 1969, Eiberle 1972). lus, Cqwa ihe.r, and S14s so-ofu) or heavily A resolution of the Swiss federal govern- reduced (Ricpicupra rupiqv-U).Along with ment approved the return of this large cat in the lynx, predators such as the wolf (Canis 1967, and Switzerland was the first country 1Lrpus). the brown bear (Ursus mctos), and to officially re-introducc the lynx into the even the scavenging (Gy- Alps in the early 1970s. This re-introduction petus harhntus) disappeared. Since the end was started as a contribution to the First Eu- of the 19th century. forests have rcgcncral- ropean Nature Conservation Year in 1970 efl, and during thc firs1 half of the 20th cen- (Lienert 1980). tury, the ungulate populations recovered The lynx population in the Swiss Alps is, quickly. This improvement inspired the idea apart from the Slovenian one. the most es- of also bringing back large predators. In the tablished of the re-founded populations, and late 1950s, reports of the expansion of the is often cited as a rare example of a suc- Carpathian lynx population towards the cessful re-introduction of a large predator 18 U. Breitenmoser et a1

(Yalden 1993). In recent years? however, Swiss Alps. A similar study was carried out the expansion of the population has halted, for the Jura Mountains (350 lynx observa- and a decrease in the number of observa- tions from 1972-87; Breitenmoser and Baet- tions in the former centre of the population tig 1992). Capt (1995) resumed this survey indicates that the population may not be as in 1992 by distributing lynx-observation viable as generally assumed. In fact, there forms to potential observers (mainly game are two lynx populations in Switzerland, wardens) in all lynx areas of Switzerland. one in the Alps and one in the Jura Moun- Fifty-thrcc direct or indirect observations tains in north-western Switzerland. In this (kills, tracks) were reported for the year report, we will focus on the Alpine popula- 1992 alone. As this method did not allow tion. The data presented and the maps discrimination between ;no report’ and ‘no shown will, however, include both popula- observation’, simplified annual question- tions, because they may have merged in naires have been sent out to all state game (Stahl and Vandel 1998). Further- wardens and hunters’ associations since more, some of our more general conclu- 1993, in order to obtain a systematic moni- sions are based on observations from the toring of the Swiss lynx populations (Capt Jura population, as in recent years field 1995). work has only been carried out in the Jura Thrcc more sources of information on the Mountains. presence of lynx are available: (1) records of livestock killed by lynx (Appendix I), (2) reports on lynx killed or found dead MONITORINGAND DATA AVAILABLE (Appendix 11). and (3) lynx trapped alive The releases of lynx in Switzerland were not for the purpose of equipping them with ra- coordinated, and there was no research or dio-collars. All domestic animals clearly monitoring programme accompanying the killed by lynx are compensated for. Up to re-introduction. None of the lynx set free in 1988, this reimbursement was paid by the Switzerland were radio-tagged. On the con- Swiss Leugue for the Protection of Nature trary, the releases were kept secret and some (SBN), and subscqucntly by the federal and official actions were even denied in public the cantonal governments. As a loss will for somc time. Consequently, there was no only be compensated for if approved by a systematic collection of observations in the qualified state game warden. these records earliest phase of re-colonization, with the are mostly an objective proof of lynx pres- exception of the examination of presumed ence, though both errors and even decep- lynx kills (Klingler and Breitenmoser 1983). tion have occurred. To instruct game war- A scientific prograinme started only in dens how to recognize lynx kills, special 1980, when problems of predation on live- courses were organized, and a brochure stock arose. Since then. an on-going rc- (Breitenmoser and Denzler 1989) and an search and monitoring prograinine has educational video (Breitenmoser et al. evolved into what we now call the Svtiss 1989) were produced. Data of lynx preda- Lyiix Project. tion on livestock have been compiled by The first study reconstructed the expansion Breitenmoser (1988), Blankenhorn et al. of the lynx population in the northern Swiss (1990), and Capt et al. (1993). The best cv- Alps, using 535 observations from the years idencc of lynx presence is a lynx found 1971-81 that were gathered from annual re- dead. The Svtiss Ljns Project maintains a ports of state game wardens as well as from data base of all known carcasses of lynx questionnaires (Breitenmoscr 1983). Haller from the two Swiss populations. Until au- ( 1992) completed the review for the central tumn 199.5. this file contained 103 records. Status of the lynx in Swikerland 19

A total of 37 lynx were caught and fitted is some uncertainty about the exact num- with radio-collars for the research projects bers and origin of the lynx set free in carried out in the northern Alps from 1983- Switzerland, because all releases were 88 (Haller and Breitenmoser 1986. Breit- made clandestinely and private actions took enmoser and Haller 1993), in the central place apart from the officially authorized Alps from 1986-89 (Haller 1992) and in operations. There was some legal uncer- the Jura Mts from 1988 to the present (Bre- tainty about the translocation of wildlife in itenmoser et al. 1993). These field studies those years. The lynx for the official. and are the main source ol' information on the probably for most of the unauthorized, land tenure system and the population dy- translocations came from the autochthonous namics of the lynx in Switzerland. population of the Slovakian Carpathian Mountains. The animals were provided by the Ostrava Zoo, who got them from hunters who captured the lynx in the wild. ORIGIN OF THE POPULKI'IONS There is, however, no proof that all the lynx The information available on the re-intro- released in Switzerland wcre really wild- duction in Switzerland has been reviewed born animals, becausc the source of some by Brcitenmoser (1983) and Haller (1992) of thc unofficially released animals is not for the Alps, by Breitenmoser and Baettig known. All lynx for the official transloca- ( 1992) for the Jura Mountains, and summa- tions were transferred to Switzerland rized by Breitenmoser and Breitenmoser- through the Base1 Zoo. where they spent Wursten (1990). At least 14 lynx were re- one month in quarantine. The animals were leased in five different places in the Swiss then brought to the site of release and were Alps, 8-10 lynx in the Jura Mts, and 3 lynx set free immediately. None of thc animals in the Swiss Plains (Table I, Fig. 1). There released was fitted with a radio-collar, and

Table 1. Releascs of lynx in Switzerland. Cantons: BE: Berne: GR: : IlJ: ; NE: Neu- chLel; OW: Obwald; VD: ; VS: . # Lynx M/F: Number of lynx (males / females) released. Fig. I: sites of relcascs as shown in Figure 1. Notes in italic indicate unconfirmed releases.

Ye'ir Site Cant on Rcgioii Fig1 # Lynx Origin Type uf re- M/F introduction

1971 Grosscs Mclchtal ow Alps 1 1/1 Ostrava official 1972 Klcinschlierental ow .4lpa 2 1/1 Ostrava official 1972 !Morrtirr- BE Jiiia b I'I 2 itriolfic.ia1 I Y72 LU Alps 2 lil Ostrava unofficial 1972 GR Alps 3 l/l Omava unofficial 1973 Pilatus LU Alps 2 iii Ostrava unofficial I974 Crew-du-Van NE Jura 6 lil O\trava official

1974 \'alldr dr. .lOlLY 1/11 Jirnr 8 2'2 7 rtrluffic~ral 197s Creux-du-Van NE Jura 6 lil Oslrava olficial 1975 CorhevriPi- 1D Alps 4 > ll~li)#/(.L7~ 197h Grand Muvcran VD Alps 4 210 Ostrava official

1976 \/n/ d'hnii~i~~s 1's flip 5 ? ril1i~f;C;d IYXO Engadiii GR 41ps 3 lil Ostrava unofficial I089 Jorat VD Plateau 9 3 1 unofficial 20 U. Breitenmoser et al

Figure 1. Sites of lynx releases in the Alps (l-S),the Jura Mountains (6-8) and on the plateau (9) oP Switzerland. 1 : Grosses Melchtal; 2: Kleinschlierental / Pilatus; 3: Engadine ( / Italian border): 4: Grand Muveran / Corbeyrier: 5: Val d'Aniviers; 6: Moutier: 7: Creux-du-Van; 8: ValICe de Joux : 9: Jorat. Further details scc Table 1. presumably only the first ones were tagged. pensation has to be covered by the cantons. Thus, the fate of the lbunder individuals re- There was some uncertainty about the ju- mained unknown. ridical requirements for translocations in the early 197Os, when the unauthorized re- leases took place. This has been clarified LEGAL STATUS with the revision of the law, as today, all Lynx was given legal protection by the translocations of protected species need to Swiss federal law in 1962. This was the ju- be sanctioned by the fede-ral authorities (JSG Art. 9l). Switzerland has signed and ridical prerequisite for the return of the large feline. With the revision of the Fe- enforced both. the Washington convention (CITES, in 1975) and the Bern Convention deral LUM,on the Hunting und Pt-otection (in 1982). which provide an international Ftve LilYrzg Miznimul,\ NII~Birds (JSG) and the corresponding decree (JSV) in 1988, the protection for lynx. protection was even strengthened. The speciea is now protected year-round (JSG Art. 7l), and a lynx causing damage to live- DEVELOPMENT A\D PRESENT STATUS OF- stock can only be shot or captured if both THE POPULATIONS cantonal and federal authorities give per- The tint ieleases in the Alps were thox in mission (JSG Art. 72,JSV Art. 10')). On the the canton of Obwald from 1971-73 (Table other hand, the confederation has to pay 30- 1. Fig. 1). Of the 2 lynx released in the SO% of all damages caused by lynx (JSG Glosses Melclrtcrl and the 6 lynx released in Art. 134, JSV Art. 14). The remaining com- the Pilatris region, an initinl population de- Status of thc lynx in Switzerland 21 veloped and spread over the northwestern part of the Swiss Alps merged with the po- part of the Swiss Alps. Because of the topo- pulation spreading from central Switzerland. graphical characteristics of this region, the The area occupied no longer increased from lynx advanced faster- towards the west than 198.5-89 (Fig. 2c), though the observations to the cast. By I98 1 - ten years after the first rose (mainly due the number of sheep killed relcascs - the population covered an area of in the canton of Valais). During the last five some 4000 km’ (Breitenmoser 1983). By years (1990-94: Fig. 2d), a reduction in thcn. the lynx released in the canton of Vaud range to the east and also a decrease in (Fig. 1) had already merged with the popu- depredations. whereas reported lynx mortal- lation of the northern Alps. Lynx also nii- ities have remained almost the same (Fig. grated into the canton of Valais (central 2). In recent years, no more observations Alps). Early observations in the cantons or were reported from areas in the cantons of Vaud and Valais. however, indicate that there Uri or Glarus (eastern Central Switzerland), were already lynx presenl - from unofficial which had continuously produced observa- releases - before the first animals from the tions in the 1980s. In the 1992/93 inquiry northern part of the Alps migrated there (S. Capt, unpubl.), the presence of lynx east (Haller 1992). Two trials in the Swiss Na- of the river Reirss was reported only from tional Park from 1972 and 1980 did not three isolated game districts. In the south- found a new population. There was never west of the population in the S any evidence of reproduction, and in spite of lynx probably migrated to the French and Schloeth’s (1978) optimistic statement that ltalian Alps. but there is no indication that the lynx was re-integrated in the Lower En- populations are established in these parts of gadine, all lynx set free in this area disap- the Alps (see also Ragni et al. 1998 and peared with fates unknown. Nevertheless, it Stahl and Vandel 1998). cannot be ruled out that the presence of lynx The expansion of lynx in the Swiss Jura in the Vdtcllina Valley () may have their Mountains until 1987 was described by Brei- origin in thc releases of 1980 at the Swiss- tennioser and Baettig (1992). Since then, the Italian border (see also Ragni et al. 1998). population has not spread any further. It is The spatial development of the Alpine po- more, likely that the area occupied has even pulation can be derived from the distribution decreased in the north-east of this (Fig. 2a-d) and the development of cases of range. The Fate of the lynx released in the depredation and of lynx killed or found dead lowlands (site 9 in Fig. 1) is not known. (Appendix I). Up to 1979 (Fig. 2a). the ob- Signs of lynx presence in this region ceased servations were somewhat concentrated in after a few observations in winter 1989/90. the region south-west of the Viei~~valdstut- Haller (1992) estiniated the lynx population tersee (Lake Lucerne) and east of Lake of the Swiss Alps at 3.5-40 adult individuals. . In the following five years ( 1980-84; This figure was based on personal experi- Fig. 2b), there was an expansion to the east ence in the northern and central Alps as well and to the west. In the east, the river Rerrss as on reported observations. Estimating the was crossed, and observations were regular- number of individuals in a large carnivore ly reported from the cantons of Uri and oc- population in forested habitat is particularly casionally Glarus. In the west, lynx crossed dilficult because the low population density the river Awe and reached the Sininzentul impedes the application of standard methods (). At least one individual such as night counts. track transects, or crossed the Gt-imsei pass (2 I65 meters) catch-and-release procedures. An approxi- from the canton of Berne towards the can- mation can be calculated if (1) the land ton of Valais (Breitenmoser 1983). During tenure system of the species is characteris- this time, the lynx released in the western tic, (2) data on individual home-range size 22 U. Breitennioser el al.

Figure 2. Distribution of the lynx in Switzcrland per five-year periods shown from the cases of preda- tion on livestock (dots) and lynx killed or found dead (stars). (a) 197 1-79: 9 domestic animals, 11 dead lynx; (b) 19XO-X4: 177 domestic animals. 23 dead lynx; following pagc: (c) 1985-89: 333 domestic an- imals. 27 dead lynx; (d) 1990-94: 185 domestic animals, 37 dead lynx (including 5 dead lynx from January - June 1995). The figures show the data reported both from the Swiss Alps and lrom the Jura Mts. The two lynx populations are separated by the Swiss Lowlands. which extend from Lake Kon- stanz to . This densely settled arid cultivated area has not been crossed by lynx so far. Status of thc lynx in Switzerland 23

C: 1985-89

are available, and (3) the total area occupied (Haller and Breitenmoser 1986, Haller is known. Lynx in the Swiss Alps occupied 1992, Breitennioser and Haller 1993). The individual home ranges from 39-425 km2 home ranges in the Jura Mts were 71-243 for females and 135-450 km' for males kin2 and 237-281 km2. respectively (Brei- 24 U. Breitenrnoser et al.

Figure 3. Distribution of lynx records in the Swiss Alps during the 1YYO-94 period. Stars indicate dead lynx (n = 36). circles livestock killcd by lynx (n = 185), and squares all other confirmed observations (n = 94). The broken line represcnts the convex polygon of the outermost observations, the solid line polygon (10,448 km2) includes the approximate area which is at present occupied by lynx. Solid lincs mark the borders of cantons and of Switzerlahd, respectively. Major lakes are shown in black. The horizontal shading represents tciniii below 600 ni, vertical shading terrain above 2000 m (the broken vertical pattern south of the polygon is a region of mainly high-alpine elevations from where digitised altitude data wcrc not available). tenmoser et al. 1993). The ranges of neigh- remaining area, 184 I km2 are agricultural bouring femalcs did not overlap (Kaczensky land, settlements or water bodies; thus we 1991), and those of males only marginally estimlztc the area at some 5000 km’ of suit- (< 10%; Diitterer 1992). We can deduce from able habitat (mainly woodland. meadows, thcsc studies that the average density in the pastures. and unproductive areas). This Alpine lynx population in Switzerland is no would conespond to 50 resident lynx for thc higher than one resident individual per 100 total area occupied in the Swiss Alps. In ad- kin2 of suitable habitat. For the 1990-94 pc- dition to these resident adult lynx, the pop- riod. a polygon drawn around the signs of ulation consists of subadult dispcrsers and lynx presence (94 observations, 185 kills of of kittens accompanying their mothers. Ob- domestic animals, and 36 lynx found dead) servations in the Jura Mts during winter re- covered an area of 10.448 km’ (Fig. 3). This vealed that juveniles added to the overall sector includes 3243 km’ of ground above density of the population by about 0.5 lynx 2000 meters and 69 1 km’ below 600 meters, per 100 km2 (Breitenmoser et al. 1993). The where lynx roam only exceptionally. Of the estimate of 50 resident lynx is still quite Status of the lynx in Swikerland 25 generous, as the polygon shown in Fig. 3 have observed a similar tendency, low sur- contains areas such as the Enrmerml- vival of young lynx did not allow the re- Nupiqehiet (about 700 km') with no obser- placement of losses among adult lynx (Bre- vations reported. Furthermore, a single ob- itcnmoser et al. 1993). The same may apply servation in Fig. 3 indicates the presence ol' for the Alpine population. However, the the lynx in the valley of the Reiiss (canton number of observations reported varies from of Uri) during the last decade. The game region to region and from year to year. wardens of this region have no longer rc- While for most areas the number of records ported any lynx observations in recent decreased, it increased in the region south- years. On the other hand, some isolated and west of the during the last three unconfirmed observations east ol' thc scttled years (Fig. 2d). Therefore, it is difficult to area (Fig. 2, Capt 1995) suggest that at least establish a general trend for the entire lynx one lynx dispersed further east. We con- population in Switzerland. clude that our density estimation is similar The stagnation 01 the population could be to Hallcr's (1992) estimate, and that there related to (1) reduccd survival of young has bccn no recent expansion of the lynx lynx and/or (2) increased mortality among population in the Swiss Alps. resident adults. Both possibilities have to be takcn into consideration. The mean age of adult lynx caught during our field study was 4.8 years (2-15 years, n=22). This rather low DISCUSSION: VERDICT ON THE PRESENT mean age for a long-living species indicates Sl'l URI'ION that the turnover - and therefore the morta- After a swift expansion towards the west in lity - among the resident lynx is high. The the first decade, the lynx population in the losses given in Appendix II are minimum Swiss Alps has stagnated over the last ten estimates. Undoubtedly, the number of uii- years, although there is still suitable habitat known cases ol lynx illegally killed is quite in the east and in the south (cantons of large. For example in the Jura Mts, only one Grisons and ). The reduced density of out of four radio-tagged lynx that were shot observations in recent years mainly in the would have been found without a transmit- east of the occupied area indicates that the ter. We assume that illegal killing is still the population has even retreated. Population most important single cause of mortality. growth was obviously not high enough to Haller (I 992) concluded that illegal shoot- encourage further expansion. Solitary terri- ing of lynx was a serious threat to the lynx torial animals such as the lynx inay not be- population in the canton of Valais. Although have very exploratively. Even during the the controversy about the lynx rc-introduc- early years after the re-introduction, the ex- tion has levelled off in recent years, and pansion to the west was probably not as first most hunter's association today formally ac- assumed by Breitenmoser (1983), bccause cept the lynx as a part of our autochthoiious the extent of the clandestine releases in the fauna, some hunters and sheep breeders still western part of the Swiss Alps had been uii- regard the lynx as a pest and - although the derestimated. Lynx remain in contact with species is protected by law in Switzerland - their conspecifics for the acquisition of high frankly state that they would shoot any lynx quality territories and subsequent reproduc- given the opportunity. One reason (or rather tion. Consequently, the capacity for expan- excuse) for the opposition of hunters to the sion of a lynx population is rather low, as lynx is the fact that the releases wcrc carried new territories are only added at the edge of out clandestinely, and some cvcn without a settled area if the population produces sur- the required permission. Though the re-in- plus individuals. In the Jura Mts, where we troduction was lengthily discussed and a 26 U. Breitenmoser et al. lasting controversy about the return of this ZUSAMMENFASSUNG predator arose, there was no follow-up pro- Der Luchs wurde in der Schweiz im 18. und gramme after the animals had been set free. 19. Jahrhundert ausgerottet. Nachdem die Those who initially released lynx believed Art 1962 gesetzlichen Schutz erhielt, be- that they wcrc best protected when lcft gann 1970 ein Wiederansiedlungsprogramm alone and no information on the project was (Tab.1). Bis 1976 wurden mindestens 14 disclosed. Although this strategy appeared Luchse aus den slowakischen Karpathen in successful in first place. it may have ampli- den Schweizer Alpen freigelassen (Abb. 1). fied and maintained the controversy. As no Eine zweite Wiederansiedlung erfolgte im authentic information was available, there Jura; zwischen den beiden Populationen was much room for rumours and the confir- besteht jedoch in der Schweiz keine mation of archaic prejudices, which are now Verbindung. Die Wiederansiedlungen wur- very hard to overcome. den am Anfang nicht uberwacht. Erst als On the other hand. we have observed low 1980 die ersten Schaden an Haustieren kitten survival in the Jura Mts (Breilenmoser (Appendix I) auftratcn, bcgann cin noch an- et al. 1993) and speculated that this inay be dauerndes Forschungsprqjekt. Die Uber- an effect of genetic problems (Breitenmoser griffe auf Haustiere erreichten 1988 niit 86 et al. 1994). There is no evidence for this hy- Tieren ihr Maximum. Seither sind sie pothesis so far. but we have to bear in mind zuriickgegangen und haben sich bei 30-40 that for the re-introductions, no precautions Tieren pro Jahr stabilisiert. Die Wiederaii- were taken to prevent genetic problems. The siedlungen in den Alpen waren unkoor- number of individuals released was low, and diniert und nicht iiberall erfolgreich. Nur die - as they presumably all originated from the Freilassungen in den Nord- und Westalpen same source - the degree of relatedness was fuhrten zur Populationsgrundung. 198 1 be- probably high. Again, the lack of infoima- siedelten die Luchse in den Schweizer tion about the translocations and the un- Alpen rund 4000 km2. Im Westen haben die known history of thc rclcascd individuals do Luchse heute Frankrcich und Italien crreicht. not allow further conclusions at this stage. Gegen Osten schritt die Ausbreitung To secure the future of the lynx populations langsamer voran und kam vor 10 Jahren in Switzerland we must establish genetic zum Stillstand. Wlhrend den letzten 5 monitoring and compare our populations Jahren beobachteten wir sogar eine Reduk- with other re-introduced or autchthonous tion der besiedelten Fliiche in1 Osten des populations. At the same time, law enforce- Verbreitungsgebiets (Abb.2). Die Luchspo- ment and education must be improved in or- pulation besiedelt heute in den Schweizer der to stop the illegal killing of lynx. Alpen eine Flache von rund 10’000 km’ (Abb. 3); wovon 50% geeignetes Luchshabitat ist. Nach der mittleren ACKNOWLEDGEMENTS. The S1.r,is.\ Ly1.r Project Wohngebietsgrosse und der Uberlappung - including the SCALP prograinme - is support- der Territorien benachbarter Luchse - ermit- ed by the Swiss Federal Office of Environment. telt aufgrund radiotelemetrischer Forcsts and (FOEFL). WWF Switzer- Uberwachungen - schatzen wir die Zahl der land, the Swiss and the Zurich Society for An- residenten Luchse im besiedelten Gebiet auf mal Protection. and the Swiss League for the ungefahr 50 Tiere. Der fur eine weiter Ex- Protection of Nature (). We thank all pansion des Siedlungsgcbicts notwcndigc the state game wardens and other people who Populationsdruck scheint gegenwh-tig zu haw rcportcd lynx obscrvations for thc monitor- fehlen. Die Zahl der iiberlebenden Jungtiere i rig programme verniag kaum die Verluste unter den adulten Status of thc lynx in Sw,itrerland 27

Tieren durch Verkehrsunfalle und illegale tial organization and recruitment of lynx Abschusse (Appendix 11) zu kompensieren. (Lynx /yn.r) in a re-introduced population in the Swiss Jura Mountains. J. Zool., London, 23 1 : 449-464. Breitenmoser, U., Rreitenmoser-Wiirsten, REFERENCES Ch. and Capt, S., 1994. Welche Chance hat der Luchs bei uns? Panda Journal, Blankenhorn, H.J., Anderegg, K.. Capt, S. 2/94: 4-5. and Brcitcnmoser. U,, 1990. 20 Jahre Capt, S., 1995. Mit langfristiger nach cincr Riickkehr in die Schweiz: Wie Uberwachung die Uberlebenschancen der gefiihrlich ist dcr Luchs den Schafen? Luchse vergrossern. Uniweltschut7- Umweltschutz in der Schwciz. BUWAL- BUWAL Bulletin, 2:38-40. Bulletin, 3/90: 21-28. Capt. S.. Bernhart. F.. Breitenmoser, U., Breitenmoser. U., 1983. Zur Wiederein- Breitenmoser-Wursten. Ch., Haller, H., burgerung und Ausbreitung des Luchses Libcrck, M., Vandel, J. M. and Herren- (Ly17.r lyt7.r L.) in der Schweiz. Schweiz. schmidt, V., 1993. Predation du lynx Z. Forstwes.. 134: 207-222. (Lynx lynx) sur Ics ongulks sauvages et Breitenmoser. U,, 1988. Luchs und Schafe. domestiques. In: Migot, P. and Ph. Stahl Dokumcntation ZLI den durch Luchse (eds). Acre du colloque “Predation ct verursachten Schiiden an Haustieren in Ckstion des Predatuers” (Dourdan 1-2 der Schweiz von 1971 bis 1988 LLI Han- Dec. 1992), Paris, 85-92. den des BUWAL und dcs SBN. Dotterer, M.. 1992. Zur Abgrenzung und Manuskript, 43 pp. Stabilitat von Territorien von Luchsmann- Breitenmoser, U. and Baettig, M., 1992. chen (Lyizx 14‘17s) im Schweizer Jura. Wiederansiedlung und Ausbreitung des Diplomarbeit Univ. Tiibingen, 7 1 pp. Luchses Lytzs 1yti.x im Schweizer Jura. Eiberle, K., 1972. Lcbcnsweise und Bedeu- Revue suisse Zool.. 99: 163-176. tung des Luchses in dcr Kulturlandschaft. Breitenmoser. U. and Breitennioser- Mammalia dcpicta, 8: 1-65, Wursten, Ch., 1990. Status, Conservation Haller, H., 1992. Zur okologic dcs Luchses Needs and Ke-introduction of the Lynx im Verlauf seiner Wiederansiedlung in Lym lyi?x in Europe. Nature and Envi- den Walliser Alpen. Mammalia depicta, ronment Series, No.45. Council of Eu- 15: 1-62. rope, Strasbourg. 43 pp. Haller, H. and Breitenmoser. U., 1986. Zur Breitenmoser, U. and Denzler, T., 1989. Raumorganisation der in den Schweizer Luchs, Fuchs, Hund - Beurteilung von Alpcn wiederangesiedelten Population Raubtier-Rissen. Wildtiere (Beilage des Luchses (Lym 1~12.1). Z. 3/11), 3:l-10. Saugetierkundc, 5 1 : 289-3 11. Breitenmoser, U., Denzler, T., Reichle, F. Hell. P.. 1961. Starkes Anwachsen der and Egli, J.. 1989. Luchs. Fuchs. Hund - Luchsbestande in del- Slowakei. Z. Beurteilung von Kaubtierrissen. Video. Saugetierkunde, 26: 57-59. 18 min. Kaczensky, P.,1991. Untersuchuiigen zur Breitenmoser, U. and Haller, H., 1993. Pat- Raumnutzung weiblicher Luchse (Lynx terns of predation by reintroduced Euro- lynx), sowie zur Abwanderung und Mor- pean lynx in the Swiss Alps. Lalitat ihrer Jungen irn Schweizer Jura. J.Wildl.Manage., 57: 144-1 54. Diplomarbcit Univ. Miinchen. 80 pp. Breitenmoser. U., Kaczensky, P.. Diitterer, Klinglcr, K. and Breitenmoser, U.. 1983. M,. Breitenmoser-Wiirsten, Ch., Capt, S., Die Identifizierung von Kaubtierrissen. Bernhart. E and Liberek, M., 1993. Spa- Schweiz. Arch. Ticrhcilk.. 125: 359-370. 28 IJ. Brcitenmoser el al

Licnert. L., 1980. Erfahrungen mit der Schmidt, W., 1960. Der Luchs in der Schau Wiedereinbiirgerung des Luchses in der von heute. Der praktische Forstwirt fur Schweiz. In: Festetics A. (ed) Der Luchs die Schweiz, 96:5 1-62. in Europa. Kilda Verlag, Grcven, 258- Schloetli, R., 197%.Luchsbeobachtungen in 263. der Schweiz. In: Wotschikowsky, U. (ed) Ragni, B.. Posseiiti, M., Mayr, S., Carrer, Der Luchs - Erhaltung und Wiederein- M.. Zangrando, E., Catello, M., Dorigat- burgerung in Europa. Bernhard, Mam- ti, E., Di Lorenzo. M., Mosca, A., Fattor. mendorf. 55-59. M. and Lombardi, G., 1998. The Lynx in Stahl. I? ‘and Vandel, J.M., 1998. Distribution of the Italian Alps. Hystrix. 10: 31-38. the lynx in the . Hystrix, 10: 3-15. Schauenberg, P., 1969. Le lynx Lyt1.r lynx Yalden. D.W., 1993. The problems of reintro- (L.) en Suisse et dans les pays voisins. ducing predators. Symp.mol.Soc.Lond., Revue suisse Zoo]., 76: 257-287. 65: 289-306. Status of thc lynx in Swit7erland 29

Appendix I: Predation on livestock by lynx in Switzerland and cornpensation paid. In the Alps, 14 cases were spccics other than sheep or goats: in the Jura Mts, 7 cases were not sheep. Other species were: 15 fallow deer (Dama dama). 1 red deer (Crn.rrs rluphus), 1 moufflon (0ii.Y ammon) (all from enclosures), and 4 cattle (calves of thc small-size “Aberdeen” breed born on pastures, unconfirmed lynx kills). Total compensation (Amount) is given in Swtiss Francs.

Year Alps Jura Mls Cases compensated

Sheep Goats Total Shccp Total Number Amount

1973 I 0 1 0 0 1 300 1975 1 0 1 0 0 1 I 50 I979 7 0 7 0 0 7 690 1980 10 1 11 0 0 11 2,470 1981 43 0 50 2 3 53 13,022 1982 44 8 53 3 3 56 17.598 1981 34 0 34 0 0 31 7.310 1984 27 0 23 0 0 23 8.890 1985 54 1 55 0 1 59 12.170 1986 57 4 61 2711 11 72 20.852 1987 32 0 42 29 73 16.29 1 1988 77 4 81 5 5 82 26,486 I989 51 12 64 8 8 72 3 I.U00 1990 34 3 40 2 2 12 13,413 1991 19 5 24 0 0 24 9.220 1992 30 2 32 0 0 32 I5.400 1991 18 3 23 2 2 25 14,930 1904 39 4 43 0 0 43 16.970

584 47 615 60 67 712 227,322 30 U. Breitenmoser et al.

Appcndix 11: Causes of lynx mortality from the two re-introduced populations in Switzerland. Spatial and temporal distribution see Fig. 2. Thc categories include: Trallic: road (23) and railway (5) acci- dents; unknown: carcasses whcrc an ananinesis was not done or no longer possible; natural: disease (4), drowning (3),or accidents (2): illegally shot: lynx shot during the hunting season or found dead with injuries cau5ed by firearms: captured: young orphaned lynx caught in the wild and put into cap- tivity: other human: other direct or indirect mortality caused by humans. The age categories are: ad: adult (>2 years); sub: subadult 1-2 years): juv: juvenile (

C'atcgnry Alps Jura Mts Total

ad sub juv total ad :\uh ~UV total

TraCflC h 3 5 I4 4 6 4 14 28 Unknohn h 2 5 15 4 0 5 11 26 Ndural I 1 I 1 2 3 1 6 9 Illegally \hot 7 2 5 17 7 0 1 10 27 Captured 0 0 5 S 0 0 1 1 6 Other human I 0 2 3 0 1 3 4 7

~ ~

Total 21 8 23 S7 17 10 IS 46 103