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Miscel.lania Zoologica 22.1 (1999) 41

Status of carnivores in (N lberian peninsula)

J. R. Aihartza, l. Zuberogoitia, E. Camacho- Verdejo & J. J. Torres

Aihartza, J. R., Zuberogoitia, l., Camacho-Verdejo, E. & Torres, J. J., 1999. Status of carnivores in Biscay (N lberian peninsula). Misc. Zool., 22.1: 41-52.

Status of carnivores in Biscay (N lberian peninsula).- The first study on the carnivore community of Biscay was published in 1985. The present study, made twelve years later, was carried out to assess the present situation of these . In this work, data collected from 1990 to 1996 by trapping, sighting, and surveying road and other kills, as well as faeces and tracks are reported. Records of 12 species were obtained, of which American Mustela vison (Schreber, 1777) is cited for the first time, and the reappears as a non-occasional species. The main trends observed during the last twelve years seem to be related to changes in land use and management of natural resources. The situation has turned favourable for the most opportunistic species, which have become very abundant in some areas, while populations of the most specialist or relictic species show a more restricted distribution. At the same time, the expansion of wolf and European mink, with the appearance of for the first time, reflect a dynamic but very unstable scene.

Key words: Carnivores, Population status, Distribution, Conservation, lberian peninsula.

(Rebut: 20 1 98; Acc. definitiva: 18 V 99)

J. R. Aihartza"), Zoologia eta Animali Zelulen Dinamika Saila, Euskal Herriko Unibertsitatea, 644 p.k., E- 48080, Bilbo, España ().- l. Z~berogoitia'~),E. Camacho-Verdejo & l. J. Torres, E. M. Icarus, C/ Pintor Sorolla 6 1" oficina 1, E-26007 Logroño, España (Spain).

O 1999 Museu de Zoologia Aihartza et al. lntrodudion along the medium and lower water courses. The vegetation was modified long The first step in biological resource surveys ago and today the lowlands are covered is to assess species richness at a determined by pine and eucalyptus plantations, and time in a particular location. Only then can the mountain tops by rough pasture. Most we proceed to a second stage, that of of the native forests have long been biodiversity monitoring, which involves cal- cleared, although some beech forests re- culating the diversity at the same location main on steep mountain slopes (LOIDI, at more than one time, in order to evalu- 1987; LOIDIet al., 1994). ate change (WILSONet al., 1996). The first During the last twelve years the area has serious attempt to assess the distribution undergone considerable changes in the en- of carnivores in Biscay was the work of vironment, mainly in relation to the pro- CASTIEN& MENDIOLA(1985). These authors gressive decline in agriculture and to the combined fieldwork with surveys carried industrial crisis of the late seventies and the out among trappers and taxidermists, data eighties. The former agricultural pattern gathered by the Natural Science Society based on small farms with cattle, fruit and Aranzadi, and previously published data. vegetables for urban consumption found it The total number of recordings and the difficult to survive. The number of cattle frequencies of each species were used to consequently dwindled and many meadows infer their (CASTIEN& were planted with eucalyptus and pine. MENDIOLA,1985). Many farms, especially those near cities, were Twelve years have since passed and dur- turned into country homes for town dwell- ing this time, land use and management of ers, and the landscape was further broken biological resources have changed consid- up by new roads and highways. At the same erably, including the establishing of sev- time, the industrial crisis sent many indus- eral Natural Parks. In this paper we com- tries into bankruptcy. As some of these en- pare data published in 1985 with the re- terprises were those which caused the great- sults of records obtained from 1990 to 1996, est amount of pollution, the environmental and changes in the community of carni- quality near urban and industrial areas im- vores are discussed. proved, and the level of pollution in some rivers decreased. The water quality was fur- ther improved as a result of several water- Study area treatment plants which were established in the area (ORIVE& RALLO, 1997). Nevertheless, Biscay (2,384 km2), located in the western canalisation of rivers continued, to the det- Basque Country (N lberian peninsula), is a riment of groves and vegetation along the densely populated territory (pop. 1,150,000), banks. with extensive urban and industrial areas. The prevailing geological materials are Cre- taceous hard limestone and flyschoid for- Methods mations, with frequent marls and sandstone. Moist NW Atlantic winds prevail. The cli- This study is mainly based on field surveys mate is consequently humid with regular carried out between 1990 and 1996. The rainfall throughout the year, ranging from data include records obtained by traps, ob- 1,000 mm on the coast to 1,900 mm in the servation of living , dead individuals mountains. A branch of the Gulf Stream, and recent skeletal remains, and identifica- with an annual thermal oscillation of 11 to tion of faeces and tracks. 12 OC, results in mild temperatures. Nocturna1 observations were made from The landscape is hilly, with altitudes 1992 in al1 the study area, travelling by car ranging up to 1,481 m a.s.1. Streams are along both tar and dirt roads. A total of short and fast, and valleys narrow and in- 64,000 km was covered over 327 nights, in- cised. Most salt marshes formed at the out- cluding al1 the 10 x 10 km UTM squares, at let of the main rivers are highly degraded, an average of 200 kmlnight. Tracking were as is the riparian vegetation, especially made by torch-light and began just after Miscel•lània Zoològica 22.1 (1999) 43 sunset, at an average speed of 40 km/h. The Results and discussion presence of carnivores was noted only when specific identification was possible. Moreo- A total of 12 species were obtained in this ver, all the carcasses found during the track- study, of which the exotic American mink ing were checked and recorded. Mustela vison (Schreber, 1777) is cited for Traps were used especially during a the first time, and the wolf reappears as a survey from 1990 to 1993. Ten live-traps were non-occasional species in Biscay. Carnivores situated in forested areas of five countries were reported in all 10 x 10 km UTM squares, (nine 10 x 10 km UTM squares): , with a mean of 4.97 species per square, and , , Gamiz-Fika and Urdaibai. a maximum of nine species (excluding occa- These traps were set for two weeks at each sional appearances, fig. 1). CASTIEN MENDIOLA point, making a total of 1900 trap-days. In (1985) reported the presence of ten species 1996, as a part of the Second Survey of carnivores in Biscay. of Spain, the main streams of Biscay were The distribution patterns of carnivores in checked for tracks following the standard Biscay are shown, comparing when possible technique by JIMÉNEZ et al. (1990). Lakes, present and past data, and their status is marshes, dams and small rivers were also sam- discussed. Distribution maps on a grid of 10 x pled in search of tracks. To avoid errors, data 10 km UTM squares are shown in figure 2. on wild and common cats, pine , stone marten, mink and polecat (see DUNSTONE, 1993), were excluded from track surveys. Family Additionally, enquiries were carried out among many colleagues surveying in Biscay, Wolf lupus (Linnaeus, 1758) as well as trappers, hunters and farmers. Records thus obtained were filtered or The wolf had disappeared completely at the checked by revision of the material (dead beginning of the century in the Basque animals, skeletal remains, photographs) Country and was still absent from Biscay in when possible. Records without reference 1985, although young were observed to be material were only accepted from qualified entering the area from the neighbouring specialists, and always after an enquiry about territory of Burgos (CASTIEN MENDIOLA, 1985). diagnostic features observed. By last winter, its presence was continuous in

VP WP uebb., mwaffina @s-,$7.3j-m®®®®1« WIUD31® ® ®:€.1 Cfl® ®Cl. rapilemen WN Fig. 1. Species richness in each 10 x 10 km UTM squares during this survey. Riqueza específica obtenida en este estudio en cada cuadrícula UTM de 10 x 10 km. 44 Aihartza et al.

Wolf Canis lupus Red vulpes

VP WP VP WP mammamamm MMOIMUGHO MODIMEffla oumoccou 0=~■M ununcccu UMEMM■■E UUDI-OccUM ■EENCE■B E■ENUM■E

European meles Pine marten Martes martes

VP WP VP WP W■■DUCO■ MMLIEMMOW EUDICIOccil ■O■M■■■E DDIMI3CODU ONMEEMEM OMMEccUM MO■M■ EMENOLIME EMENEMB

Stone marten Martes foina Mustela erminea VP ■■■■E■ WP VP WP MMINU3030. 3MNMMMEffl ■LII■M■E OMMER31.110U UOMEMIMIU Offin0HccDel UffiroMMIOMM MENNOLME E■ENZE■E

Fig. 2. Distribution maps of different species: white circles indicate presence in the square before 1986 (bibliographic data); black circles, presence after this year (new data); divided circle, presence before and after 1986; asterisk, the occasional appearance of the wolf. Miscel.lania Zoologica 22.1 (1999) 45

Weasel Mustela nivalis European mink Mustela lutreola

American mink Mustela vison Western polecat Mustela putorius

Wild cat silvestris Common genet Genetta genetta Aihartza et al. the western tip of our study area. Between verse food, and where the risk from hunters 1988 and 1992, farmers shot ten in and illegal trappers is low. Karrantza and Artzentales following attacks The previous explanations in accounting on livestock. One hundred and six sheep, five for the return of the wolf are also applicable goats, five calves and six young horses were in the trends of other carnivores, the killed (Saenz de Buruaga, Campos & Onrubia, in particular. Moreover, the progressive aban- pers. com.). This carnivore was present dur- donment of farmlands in Biscay with substi- ing the summer from 1994 to 1997 and a tution of croplands by pine or eucalyptus bitch with breeding signs was killed in plantations, and the consequent spreading Artzentales in summer 1994. Breeding in the of scrub and bushland causes an increase in area was confirmed in spring 1997 when a potential roosts. The expansion of red fox in cub was killed in (E. Álvarez, pers. com.). Scotland was also correlated with an increase The increasing presence of wolves in Bis- in potential roosts (HEWSON& KOLB, 1973). At cay reflects its general expansion in Europe the same time, there has been an increase in and in particular its spread towards the south "security areas" surrounding towns and cit- and east of the lberian peninsula (BLANCOet ies, where hunting is strictly forbidden, and al., 1990; BLANCO& GONZÁLEZ,1992; PROMBERGER where many small and medium-sized carni- & SCHRODER,1993). The main reasons for the vores roam in safety. Nevertheless, its present expansion of wolves in the lberian peninsula abundance is threatened by sarcoptic mange seem related to changes in land use and (LINDSTROMet al, 1995), which is decimating conservation policy (BLANCO,1997). On one populations of in neighbouring areas hand, the progressive abandonment of farm- of western Pyrenees and Basque Country, lands has allowed natural forests and shrubs and has recently been detected in southern to spread, and resulted in an increase of wild Biscay where eight dead foxes were found boar Susscrofa (Linnaeus, 1758) and roe deer and at least 14 sick foxes were seen. Capreolus capreolus (Linnaeus, 1758). This trend is clear in Biscay, where the migration of wolves from the west and south west coin- Family cides with that of wild boar and roe deer. On the other hand, prohibition of poison use in European badger Meles meles (Linnaeus, 1983, and increased conservation measures 1758) favoured the return of this canid. Present data agree with the classification of CASTIEN & MENDIOLA(1985) as a widespread Red fox Vulpes vulpes (Linnaeus, 1758) species in Biscay. Although less frequent than other carnivores, it seems to have spread dur- The red fox, a widespread in the ing the last decade. The badger is distributed lberian peninsula (CASTELLS& MAYO, 1993), throughout the territory, but few contacts was reported to be the most abundant car- per square were obtained during the present nivore in the western Basque Country in study. It is found in a great diversity of habi- 1985 (CASTIEN& MENDIOLA, 1985). Present data tats in the province, with preference for coun- even suggests an increase in frequency, since tryside and fields with scattered thick forest foxes have been in al1 the squares and al1 patches. The badger also seems favoured by kinds of habitats. In 1991, fox densities in the increase of thick forests due to the ex- central and eastern Biscay were estimated to pansion of pine plantations. reach 6.75 ind./km2, which is one of the Nevertheless, its real status is difficult to highest values recorded in Europe (REIJAet determine because of the lack of specific al., 1991). Thus, the red fox can be consid- research. GRIFFITHS& THOMAS(1993) noted ered even as an "overabundant" species in populations and trends of this species in the some areas (WAGNER& SEAL, 1992; GARROTet lberian peninsula remained unknown. Its al., 1993). In the present survey, red fox seems presence was reported to be well established to be more frequent in lowlands, outskirts in Catalunya (Ruiz-OLMO,1990), but it seems of small villages, and especially near rubbish to be threatened in the central lberian pe- dumps, where it can find abundant and di- ninsula (VIRGOS,1994). Miscel.lania Zoologica 22.1 (1999)

Pine marten Martes martes (Linnaeus, 1758) Natural Parks) where this species seems to be very common. These data agree with the The pine marten is a rare carnivore in Biscay previous work of CASTEIN& MENDIOLA (1985). area and its distribution was reported to be limited to the oak and beech forests in the mountains of the south and west (CASTIEN& Stoat Mustela erminea (Linnaeus, 1758) MENDIOLA,1985). Five pine were lo- cated during the present study in two squares This seems to be a rare carnivore in Biscay. of the western tip of Biscay: in Karrantza CASTIEN& MENDIOLA(1985) considered it to be and Turtzioz three individuals were observed restricted to the occidental area of Biscay. foraging, a fourth (a cub) was captured by a Nevertheless, although al1 our records (12 farmer and left in the rehabilitation sightings plus three carcases) were obtained center "El Carpín", in Karrantza, and an- in four squares of the western tip of the other was found dead. No individuals were study area (Karrantza and Turtzioz), it was oberved however in the southern moun- recently recorded by A. Galarza (pers. com.) tains of the study area (squares WN16 and and G. Ocio (ZUBEROGOITIAet al., 1997) in WN26), where it was recorded by CASTIEN& Gorbea Natural Park (S Biscay). It seems un- MENDIOLA(1985). likely that the lack of previous recordings Although hunting of small carnivores is could be accounted for by scarce surveying, now illegal, and pressure on pine marten as this species was highly valuable for trap- has consequently been reduced, there has pers and hunters. The stoat therefore may not been any noticeable increase in the be spreading in Biscay throughout the moun- populations of this mustelid, and it seems it tains ranging eastward from Karrantza to has disappeared from some areas. Other en- Gorbea. vironmental factors must therefore be in- volved. There is considerable controversy about the feeding of pine marten (ERLINGE, Mustela nivalis (Linnaeus, 1766) 1986; THOMPSON& COLGAN, 1987). Recent stud- ies in the Cantabrian Mountains showed that were found in sixteen squares, in al1 the frequency of woodland rodents is high kind of habitats from sea level up to 1,000 m in the pine marten's diet, suggesting that a.s.l., showing preference for countryside. this mustelid forages primarily in beech or However, its frequency is lower than that of oak forests, and less frequently in open habi- species such as genet, stone marten and tats (CLEVENGER,1993). The same author noted badger. The weasel was reported as the most that its diet varies according to food avail- frequent carnivore (CASTIEN& MENDIOLA, 1985), ability and degree of competition. Thus, the and it was recorded in fourteen squares in fragmentation and scarcity of deciduous for- Biscay. It has been considered a widespread ests could be a significant handicap for the non-threatened species in the lberian pe- expansion of pine marten, particularly if high ninsula (CASTELLS& MAYO, 1993). Nevertheless, densities of opportunistic carnivores (such as there has been a decline in records through- foxes, stone martens or even genets) exert out the study area in recent years. This is strong competitive pressure in surrounding surprising since pressure against small carni- areas (BRAINERD,1997). vores has decreased and habitat availability appears unchanged. Furthermore, there is no evidence of rodent poisoning and other Stone marten Martes foina (Erxleben, 1777) rodent predators such as pine marten, fox, barn Tyto alba (Scopoli, 1769) or tawny According to our data, the stone rnarten is owl Strix aluco (Linnaeus, 1758) are abun- one of the most frequent carnivores in Bis- dant (ZUBEROGOITIA& TORRES, 1997). The rea- cay. It was located in al1 kind of habitats son for the decline in weasel populations from seashore up to Gorbea Mountain (1,400 therefore remains unknown but may be due m a.s.1.). Records were most frequent in the to competition or predation. Although com- western tip (Las Encartaciones) and the petition of stoat with weasel has been docu- southern mountains (Gorbea and Urkiola mented (KING,1989), the stoat is restricted 48 Aihartza et al. only to a small area. Raptors do prey on American mink Mustela vison(Schreber, 1777) weasel (MIKKOLA,1983) however, and during the present work three cases of predation by The first records in the lberian peninsula domestic cats were recorded, as well as one were reported by DELIBES& AMORES(1978) by the barn owl and one by the tawny owl. and it was later observed in the Central Sys- Although there are no quantitative data to tem DELIBES(1983). In 1987, VIDAL-FIGUEROA& assess the significance of predation by this DELIBES(1987) confirmed its presence in SW species, it must be noted that the tawny owl, Galicia and in NW Portugal. RUIZ-OLMO(1987) with more than 1,700 couples located in reported its distribution in diverse regions in Biscay, presents one of the highest densities Catalunya. BUENO& BRAVO(1990) also noted in Europe (ZUBEROGOITI~& TORRES, 1997). quick expansion throughout the Central Sys- tem. Finally, Ruiz-OLMOet al. (1997) extended the area cited to and Teruel- European mink Mustela lutreola (Linnaeus, Castelló. 1761) The American mink was first observed in Biscay in 1993 in the Butroe River, when Fourteen records of European mink were some people saw "otter-like" animals. Two obtained in Biscay: eight observed alive, and years later, the authors captured an Ameri- six found dead. Most data were obtained in can mink at this site. More data were later medium and lower river courses in the north, collected from the same and other nearby and the rest in the southern mountains as rivers such as the Oka and the Asua (three well, in Gorbea and Urkiola Natural Parks. individuals were trapped and 15 direct ob- The present distribution includes the rivers servations made). Likewise, another indi- Artibai, Ea, Oka, , Butroe, Nerbioi, vidual was trapped in the Nervion river in lbaizabal and Urkiola, ranging from sea level September 1996 by County Council rangers, up to 700 m a.s.1. Although several authors and taken to the animal rehabilitation (e.g. BLANCO& GONZALEZ,1992) stated that center, El Carpín, in Karrantza. are linked to well conserved streams, This population proceeded from escapes in Biscay they can be found even in moder- and releases over the last decade, when many ately polluted and perturbed streams. permits for fur farms were issued in the study Commercially valuable fur-bearers are area: 18 fur farms were recorded in the Au- amongst the first species to be recognised tonomous Community of the Basque Coun- by local hunters, trappers and naturalists try (where Biscay is included) in 1992, with (YOUNGMAN,1982). Nevertheless, the Euro- an estimated total stock of 10,000 females pean mink was first reported in the lberian (PALAZ~Net al., 199813). peninsula in 1951 (RODRIGUEZDE ONDARRA, An estimate of feral population and dis- 1955). It appears likely that this species spread tribution is required to know its present sta- into the Basque Country from coastal tus and prevent possible problems: competi- during the mid-1940's. as a result of high tion with other carnivores such as stone mar- population density, according to the many ten, polecat or the European mink, and sexual individuals preserved in museums at that competition with the latter, as suggested by time (YOUNGMAN,1982; PALAZÓN & Ruiz-OLMO, some authors, although hybridisation does 1992; Ruiz-OLMO& PALAZ~N,1995). During the not seem to occur in nature (YOUNGMAN,1982; following years many data were obtained in MARAN& HENTTONEN,1995). the Basque Country and neighbouring re- gions, confirming its expansion (RODR~GUEZDE ONDARRA,1963; BLASARITIO, 1970; SENOSIAIN & Western polecat Mustela putorius (Linnaeus, DONAZAR,1983). 1758) Although there are no data on popula- tion densities, the European mink seems to The polecat was recorded at 12 sites during continue to be increasing both in distribu- this study: , , Abanto and La tion and frequency in nearby areas such as Arboleda, a mountain coniferous forest in Araba, Navarra, La Rioja and Burgos (TORRES Karrantza, and in the streams Aguera, & ZUBEROGOITIA,1996; PALAZ~Net al., 1998a). Barbadun, Kadagua, Butroe, Lea and Miscel.lania Zoologica 22.1 (1999) 49

Karrantza. These data include two road kills, European countries (HUBBARDet al., 1991; the shooting of a family and nine sightings. DOTTERER& BERNHART, 1996). Some of these records were in highly popu- lated areas. CASTIEN& MENDIOLA (1985) recorded the polecat as a widespread but infrequent Family species. These authors noted that the scarcity of polecat in the western Basque Country Common genet Genetta genetta (Linnaeus, could be due to the little value of its fur. 1758) Nevertheless, our data (in which trappers and hunters playa minimal role) confirm its wide- This is one of the most common carnivores spread but infrequent status. lnterpretation is in Biscay. It was found frequently through- somewhat difficult as ecology and behaviour out the study area, in al1 kind of habitats, of the polecat is poorly understood elsewhere ranging from sea level, in Urdaibai marsh, (BLANDFORD,1987). was to 1,100 m a.s.1. in Gorbea Mountain. It was pointed out as an important factor in the noted also as a widespread carnivore in the decline of the polecat on the continent. Some western Basque Country (CASTIEN& MENDIOLA, authors considered that drainage of ponds 1985), and in the lberian peninsula as well and marshes, canalisation of rivers and deg- (CASTELLS& MAYO, 1993). radation of wetlands, as well as the destruc- tion of deciduous woodland limited the habi- tat availability for this species in different General discussion countries (LODE,1995). Our data agree with this viewpoint since the polecat occurs mainly The community of carnivores in Biscay shows in the best preserved areas and streams in a pattern typical of industrial and densely Biscay. The scarcity of deciduous woodland populated areas in SW Europe. The big car- and well-conserved river corridors and nivores such as brown ( arctos, wetlands in this area also account for its low Linnaeus 1758), wolf [Canis lupus (Linnaeus, frequency. 1758)] and (Lynxsp.) disappeared many decades ago, mainly as a result of direct hunting and loss of habitat (BOUCHET,1988; Family KEMPF,1987). Small and medium-sized fur- bearers such as the otter ( lutra, Linnaeus Wild cat Felis silvestris (Schreber, 1777) 1758) disappeared in the same way or be- came reduced to residual populations (DELIBES, Eleven records of wild cats were collected 1990). during the present study, including two The highest species richness was recorded trapped, two road kills, and seven direct in Karrantza and Turtzioz, in the western tip observations including one occupied bur- (VN67, VN68, VN78 and VN79) with eight or row. All were located in the best conserved nine species. All these squares have a high mountain areas of the west and the south. diversity of habitats, including pristine de- This agrees with CASTIEN& MENDIOLA(1985), ciduous forests and well-conserved streams. who ;ssociated the presence of wild cats These characteristics allow the presence of with the best preserved forests in the west- the most specialist or residual species such as ern Basque Country. the pine marten, wjld cat, stoat or polecat. Little is known about this carnivore in Excepting the wolf, which returned only re- the lberian peninsula, apart from its wide- cently, these species were recorded in the spread distribution (CASTELLS& MAYO, 1993). previous work of CASTIEN& MENDIOLA(1985), Habitat loss was noted as one of main threats though not in al1 squares. both in Spain and in other countries in the Excluding marginal squares, the lowest European Community (GINN,1984; BLANCO& richness appears today in some areas of cen- GONZALEZ,1992) while recent studies have re- tral Biscay with three or four species (red ported that hybridisation with feral cats is fox, genet, stone marten or badger). In these one of the main reasons to explain the ex- squares extensive areas are covered by pine tinction of wild cat populations in some and eucalyptus plantations alone, valleys are 50 Aihartza et al. highly developed and streams degraded res, e identificación de excrementos y huellas. (ORIVE& RALLO,1997). It is important to ana- Se han obtenidos datos de 12 especies, entre lyse highly populated areas near big cities, las cuales el visón americano Mustela visonse such as squares VN99, WN09, WN19 and cita por vez primera en Bizkaia, y el lobo WPOO, where seven species were recorded: reaparece como especie no ocasional the most opportunistic and widespread spe- (figs. 1, 2). Las principales tendencias obser- cies such as red fox, stone marten, genet or vadas están relacionadas con cambios en el badger, plus weasel and two species of minks. uso de la tierra y gestión de los recursos Finally, comparing data with the previous naturales. Las especies más oportunistas han work of CASTIEN& MENDIOLA(1985), and keep- incrementado su presencia llegando a ser muy ing in mind that our work has a higher abundantes en algunas áreas, mientras que sampling effort, a decline in species richness las especies más especialistas o con poblacio- can be observed in squares WN26, WN29 nes relictas se hallan estancadas o en declive. and WN49, from eight, seven and seven spe- Al mismo tiempo, la expansión del lobo y del cies respectively, to six, five and five. This visón europeo, junto con el asilvestramiento variation is due to the absence of records del visón americano, plantean un escenario about pine marten, polecat and weasel dur- dinámico aunque muy inestable. ing the first work. Possible explanations are given above for each of these species. In conclusion, the main changes observed Acknowledgements in the conservation status of carnivores over the last twelve years seem related to changes We wish to thank al1 those who took part in land use and management of natural in the field work, especially Esther Bernedo, resources, including conservation efforts and Luisa Fernanda Campos, Eneko Imaz, Mari attitudes of hunters and farmers. Neverthe- Jose Totorika, Julen Zuberogoitia. We also less, these factors do not affect al1 species appreciate the collaboration of those col- in the same way. In short, the most oppor- leagues who provided valuable data: Pedro tunistic species such as red fox, stone mar- Abad, Lander Astorkia, Arturo Elosegi, Aitor ten, genet and even badger are frequent Galarza, Fernando Garitagoitia, Jose Antonio or very frequent. At the same time, González-Oreja, Javier Gómez-Abrisketa, Jose populations of the most residual species Antonio Mugica, Fernando Ruiz-Moneo, seem to have declined, ¡.e. weasel, polecat Edorta Unamuno, Aitor Uriarte. Our sincere pine marten and wild cat. This may be re- appreciation to Drs. E. Castien, A. Elosegi, lated to limited habitat availability or l. Garin, l. Mendiola and J. Ruiz-Olmo for interspecific relations. Other reasons such as their critica1 review of the manuscript. pollution, disease, or hunting may also play a role, although there is no consistent evi- dence of this. References

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