distribution of the iberian wolf in galicia. concordance between field sampling and questionnaires....

7/27/2019 Distribution of the Iberian Wolf in Galicia. Concordance Between Field Sampling and Questionnaires. L. Llaneza & P

1/10

ORIGINAL PAPER

Wildl. Biol. Pract., June 5(1): 23-32DOI: 10.2461/wbp.2009.5.5

Distributionofthe iberian Wolf (Canislupussignatus) in Galicia (nW spain):

concorDance betWeen fielD samplinGanD Questionnaires

L. Llaneza* & P. Nez-Quirs

A.RE.NA. Asesores en Recursos Naturales, S.L., c/ Perpetuo Socorro, n 12 Entresuelo 2-B 27003 Lugo,

Spain

* corresponding author: e-mail: [email protected]

Keywords

Wolf,

Distribution,

Questionnaires,

Sampling methods,

NW Spain.

Abstract

The information about wolf presence in Galicia (NW Spain) obtained

from questionnaires sent to wildlife rangers (163 with a 59.50% response)

and huntig associations (629 with a 28.29% response) is compared to

that obtained from eld sampling involving the surveying of 305 UTM

10x10 km squares. The data were recorded as presence/non-presence in

UTM 10x10 km squares. Wildlife rangers provided information on wolf

presence in 58.03% of squares, Hunters Associations in 52.13% and eld

sampling in 63.60%. The three sources of information differed with respect

to the number of grids with wolf presence/non-presence. The overall

percentage of concordance (presence/non-presence) between sampling

and questionnaires was 70.16%. Concordance between eld sampling and

questionnaires was 85,57% for wolf presence and only 43,24% for wolf

non-presence. Although the present results suggest that questionnairesprovide reliable information in areas where wolves have usually been

present, they may leed to overestimation of wolf presence on humanised

land and underestimation in areas near provincial boundaries.

Introduction

Knowledge of the distribution area of species is a fundamental tool in studies focusing

on the conservation and management of the natural heritage [1]. Carnivore distributionis hard to establish [1,2] since carnivores are (1) difcult to observe and identify inthe eld, (2) evasive, (3) often nocturnal and (4) found only at low densities. Thus,researchers are obliged to locate tracks, scats and food remains in order to ascertain

their distribution range [3]. Field sampling can be feasible in small areas [2], but forlarge-scale studies where little is known about the biology of a particular species,questionnaire surveys among a variety of social groups may be more suitable [4] ,especially for widely distributed species [5].Questionnaires are often used as a source of information on the distribution [6-11]and population trends [2,12] of several species, especially carnivores, being used

particularly to collect data on hunting activities [13,14]. As regards wolves, they have

been used in local studies [15-18] and large-scale studies [19-21].On the Iberian Peninsula, the iberian wolf (Canis lupus signatus) and Man havelived in conict for millennia due to wolf depredation on livestock and competitionfor game. Thus, Man has acquired a sound knowledge of wolves and their breedingsites with a view to eradicating them. Certain social groups may,therefore, be able to

provide useful information on wolf presence and distribution.
http://dx.doi.org/10.2461/wbp.2009.5.5http://dx.doi.org/10.2461/wbp.2009.5.5http://dx.doi.org/10.2461/wbp.2009.5.5


2/10

24

Galicia, in the NW of the Iberian Peninsula, has a long history of wolf presence[22-23]. Wolves generally occur in areas with a high density of human populationand numerous infrastructures. The Galician landscape is characterised by uniform

distribution of small settlements (villages, hamlets, farms, etc.). The lack of largetracts of uninhabited land inevitably leads to the close coexistence of wolves and

people. In addition, abundant available livestock contributes to the otherwise unusualformation of wolf packs in highly humanised areas [16]. Consequently, certaingroups, such as shepherds, cattle breeders, nature managers, wildlife rangers and

members of hunting associations are very aware of wolf presence. The latter two canprovide information on wolves due to their work or recreational activities.Although questionnaires have been used in various studies on wolves in the IberianPeninsula, none has assessed and compared data thus obtained with data collected by

researchers in the eld. In this study, we compared data on wolf presence in Galiciagathered from three different sources: questionnaire responses from wildlife rangers,responses from hunting associations and the results of eld sampling. The usefulnessof questionnaires as a method for establishing the large-scale distribution range of

wolves in Galicia was assessed.

Material and methods

This study is part of broader research into the distribution and population status of theIberian wolf in Galicia, conducted between 1999 and 2003 [16-18]. During that period,the three main methods used to collect data on wolf presence were: questionnairessent to wildlife rangers (n = 163), questionnaires sent to hunting associations (n = 629)

and eld sampling by our research team. Questionnaire design and forwardingfollowed the proposals of Filion [24]. Firstly, a letter was sent to each group settingout the survey objective [7], emphasising the importance of collaboration andexplaining the format of the questionnaire that they would be asked to ll in andreturn. Simple short questionnaires basically consisting of concisely written yes-noquestions requiring short answers were then sent, accompanied by clear and briefinstructions. Pre-paid postage envelopes were included. The information requested

referred to sightings, mortality, livestock damage and reproduction. On the basis ofthe questionnaires, data on wolf presence were mapped on to UTM 10x10 km grids.In the squares for which the two groups of informants provided no information, weassumed non-presence of wolves. The data thus gathered were carefully analysedfor mistakes and anomalous or meaningless data (births outside the normal period:

April-June, etc.), which were immediately rejected. The territorial distribution of theinformants according to working areas (wildlife rangers) and game reserves (huntingassociations) was homogeneous in the study area.Field sampling conducted by our research team covered the whole of Galicia, exceptfor the very developed coastal strip, highly populated areas and places where wolf

presence cannot be conrmed. Overall, the research team covered 305 of the 365UTM 10 x 10 km squares (Fig. 1). Field sampling consisted of transects performed inhabitats within each 10 x 10 km square with the highest probabilities of containing

wolf signs (mainly scat and scratch marks), i.e. dirt roads, rebreaks and paths [25],thus avoiding the bias of sign detection that occurs in habitats with varying degreesof vegetation cover. Transects were surveyed on foot or in all-terrain vehicles, that


3/10

25

enable large areas to be covered at low cost.Only the 305 UTM squares sampled by our research team were compared with thedata obtained from questionnaires. In this paper, besides wolf presence we will

also refer to non-presence in acknowledgement of the fact that both eld samplingand questionnaires may fail to detect presence in a given UTM square. Sixty UTMsquares considered unsuitable for wolves were not surveyed, being scored as showingnon-presence.

We have assumed eld sampling as a reference baseline and comparing it withquestionnaires to assess their relative performance, we do not intend to imply thateld sampling is superior, only that the more time-consuming method (eld sampling)is used as a reference against which to compare the more economical method of

questionnaires

The G-test was used to compare the frequency of presence/non-presence among thethree sources. In addition, concordance between eld sampling and the other twodata sets was tested by logistic regression. For this purpose, eld sampling was usedas the dependent variable and the questionnaire data were the independent variables.The advantage of logistic regression, compared to the commonly utilised G-test, is

Wildlife Rangers Hunters Associations

Presence

Non presence

Non sampled grid

Field sampling

Fig. 1. Grids showing presence-non presence of wolves according to the different sources of information.


4/10

26

that it permits consideration not only of overall agreement, but also of agreementin presence (specicity) and non-presence (sensitivity) [26]. This is interesting

because disagreements between data sources can be due to differential performance

in recording presence (differences in specicity) or in differential performancein recording non-presence (differences in sensitivity). Thus, this information isrelevant to assessing the merits of each sampling strategy. Although eld samplingwas considered as the response variable in this analysis, the issue of which variableis considered as the predictor and which the response variable is not essencial inapplying logistic regression to this data set.The degree of concordance between eld sampling and questionnaires may varydepending on wolf density (detectability) and local peoples experience with wolves(length of historical presence). We tested this possibility by dening three zones in

Galicia: (1) those where wolves have historically been present equivalent to zoneswith high wolf density; (2) transition zones equivalent to areas of average density;and (3) highly humanised zones equivalent to areas where wolves are absent oroccur at very low density (Fig. 2).

Results

Ninety seven questionnaires (59.50%; Table 1) were returned by wildlife rangers,providing information on wolf presence in 177 squares (58.03% of total squaressampled by our research team) and no indication of presence in 128 squares (41.97%)(Fig. 1). From hunting associations we received 178 questionnaires (28.29%; Table1), indicating wolf presence in 159 grids (52.13%) and suggesting non-presence in

146 squares (47.87%) (Fig. 1). Field sampling indicated wolf presence in 194 grids(63.60%) and non-presence in 111 (36.39%) (Fig. 1).Results indicate that 70.15% of squares (54.42% for presence and 15.73% for non-

presence) showed agreement between eld sampling and questionnaires (Table 2;Fig. 3). In only a small percentage of squares did eld sampling indicate presence andquestionnaires indicate non-presence (9.18%; Table 2; Fig. 4).

Fig. 2. Map representing the three geographical areas in which Galicia can be divided according to wolf

presence.

Highly humanized zone

Transition zone

Zone with historical presenceof wolves


5/10

27

Table 2. Survey team and questionnaires coinciding regarding presence (1) / non-presence (0).

Coincidence

Survey Team Questionnaires Number of grids %1 1 166 54.42

1 0 28 9.18

0 1 63 20.65

0 0 48 15.73

Table 1. Results of questionnaires sent to wildlife rangers and hunting associations in the provinces ofGalicia.

Provinces Wildlife Rangers Hunters Associations

Number ofquestionnaires Number ofanswers % of answers Number ofquestionnaires Number ofanswers % of answers

Pontevedra 33 20 60.60 70 31 44.28A Corua 40 17 42.50 132 44 33.33

Lugo 50 30 60 260 64 24.61

Ourense 40 30 75 167 39 23.35

Total 163 97 59.50 629 178 28.29

Fig. 3. Coincidences in presence/non-presence of wolves between eld sampling and questionnaires.

Coincidence

Non coincidence

Non sampled grids

Fig. 4. Grids in which eld sampling denotes presence and questionnaires indicate non-presence ofwolves.

Survey team presence,questionnaires non presence

Rest of sampled grids

Non sampled grids


6/10

28

The different sources of information differed signicantly in the number of squareswith presence/non-presence of wolves (G = 8.26; d.f. = 2; p = 0.016). There were nodifferences in the number of squares with presence/non-presence of wolves between

eld sampling and wildlife rangers (G = 1.989; d.f. = 1; p = 0.158); however asignicant difference existed between eld sampling together with wildlife rangersversus hunting associations (G = 6.20; d.f. = 1; p = 0.012).The logistic regression comparing the concordance between eld sampling andquestionnaires (wildlife rangers plus hunting associations) indicated signicantagreement between both data sets. The concordance table of the logistic regression

showed that agreement with eld sampling was 85.57% (specicity) for the squaresin which the questionnaires indicated wolf presence and 43.24% (sensitivity) for thesquares in which questionnaires indicated non-presence (p < 0.001).

There were differences between the three zones with respect to the number of squares

with wolf presence (G = 39.774; d.f. = 2; p < 0.001). There were no differences in thenumber of grids with wolf presence (G = 1.736; d.f. = 1; p = 0.188) between zones1 and 2 but there were differences between zones 1 and 2 with respect to zone 3

(G = 38.03; d.f. = 1; p < 0.001). Some of the squares with presence mentioned in thequestionnaires were bound to certain geographic areas. Thus, in the central-western

zone (Zone 3), eld sampling indicated wolf non-presence whereas questionnairesshowed presence (Fig. 5). In Zone 3, questionnaires and eld sampling showedconcordance in wolf presence for 27.11% of squares, whereas in Zone 1 concordance

was 64.70%.

Discussion

Although questionnaires are commonly used to collect information about variousaspects of animal species (distribution, population trends, hunting activities, etc.),the data that they yield need to be interpreted with caution [27] because of thevariety of sources and varied experience of the informants [12]. Furthermore, thedistribution of the people surveyed must be representative of the study area [5], acondition fullled by our data. Data reliability also needs to be checked and dubiousinformation discarded, both in questionnaires and interviews [11,28,29]. We also

Fig. 5. Grids in which eld sampling denotes non-presence whereas questionnaires indicate wolfpresence.

Survey team non presence,questionnaires presence

Rest of sampled grids

Non sampled grids


7/10

29

took this into account. Other factors may cause wolf presence to be underestimated oroverestimated., e.g. lack of experience in animal observations can lead to confusion,and there may be political reasons for some people being interested in inaccurate

estimates of carnivore presence for political reasons [30]. This may occur in the caseof wolves as a result of the social conict they generate. Many people (chiey farmersand hunters) are interested in demonstrating that wolf numbers have increased inorder to call for strict culling policies. Others (e.g. environmental groups), on theother hand, may wish to prove numbers have fallen so as to prevent eradication.Many authors recommend robust sampling designs and questionnaires to reduce theimpact of such misinformation [5]. If carefully planned, interpreted and controlled,questionnaires can yield valuable results [30]. Other authors combine questionnairesand eld interviews to check the information obtained, an approach that reduces the

disadvantages of using each method separately [31] and increases efciency [10].Moreover, research studies such as this one permit the validation of this method ofascertaining distribution ranges as compared with eld sampling. Despite the abovementioned caveats, we agree with other authors that questionnaires are a valuableway of collecting data in general [2,5,6,8,11], and, in particular, information onthe distribution of Iberian wolves in Galicia. However, there are some problemsassociated with information obtained via questionnaires. The questionnaires wereinitially designed as a starting point for future studies [8,11,30].Both wildlife rangers and hunting associations provided similar levels of information;in fact, no signicant differences were found between the two sources regarding thenumber of squares showing presence/non-presence. This may be due to the regularcontact between both groups and mutual inuence due to the sharing of informationabout signs, direct sightings, wolves being heard or damage to livestock. Nevertheless,hunters indicated the lowest number of squares with presence and yielded the largestdifference compared with eld sampling data. These divergences could be due to thelow response rate in this group [4]. It is striking that in areas where eld samplingindicated non-presence, the information from questionnaires showed presence (Fig.5), especially in the highly humanised areas (Fig. 2). This leads us to believe that theinformation on presence in the questionnaires may represent an overestimation of

wolf presence. The questionnaire data from that area refer mainly to livestock damageattributed to wolves. Such information on damages caused by wolves must be takenwith caution since it is well known that damage caused by dogs, false claims bycattle breeders aiming to obtain compensation and unconrmed rumours of livestockdamage are common in the wolf distribution range on the Iberian Peninsula.

The squares for which eld sampling indicated wolf presence and questionnairesindicated non-presence show a clear spatial pattern, representing either zones near

the coast or provincial boundaries. In the former, the people surveyed may barely beaware of the presence of wolves. Close to provincial boundaries, people may have

failed to detect wolf presence because the square was partially located outside thelimits of their working areas (wildlife rangers) or their hunting reserves (hunters)(Fig. 4). Therefore, information on wolf presence in peripheral zones may beunderestimated. Such situations can occur particularly in zones where wolves havenot been present in the last 40-50 years and, as a result, inhabitants do not recall them.The fact that no data exist on carnivore presence in a certain area does not mean that


8/10

30

there are none there [6,30]. Interviews, for example, have been showed only to besuitable for areas often frequented by the interviewees [32].On the Iberian Peninsula, wolf studies have involved the partial use of questionnaires

[20,21,33] in order to establish distribution range and to collect complementaryinformation. They have not been used directly to determine wolf density. A study

based on questionnaire responses from hunters to assess wolf density showed anoverestimation of real density. In fact, most observations were concentrated in areaswith a high density of prey, whereas in other zones with no game, wolves were notdetected [34].We consider that questionnaires sent to rangers and hunters can contribute information

to determine wolf distribution ranges. In zones where wolves have historicallyoccurred and in zones with high wolf density, information from questionnaires can be

considered reliable. However, in humanised zones with no historical wolf presence orwhere wolves occur at low density, questionnaires may overestimate wolf presence.

Acknowledgements

Special thanks to Wildlife Rangers of Xunta of Galicia and Hunters Associations of Galicia. Andrs

Ordiz, Antonio Uzal, Vicente Palacios, Vctor Sazatornil, Pablo Sierra, Francisco Alvares, Pedro Alonso

and Jorge Layna carried out the eld work. Marcos Mndez made valuable comments to a previous draft

of this paper.

References

1. Palomo, L. J. & Gisbert, J. 2002. Atlas de mamferos terrestres de Espaa. [Spain mammals atlas]

Direccin general de conservacin de la Naturaleza-SECEM-SECEMU. 564pp.

2. Lovell, C.D. Leopold, B.D. & Shropshire, C.C. 1998. Trends in Mississippi predator populations,

1980-1995.Wildl. Soc. Bull. 26: 552-556.

3. Zielinski, W.J. & Kucera, T.E. 1995. American Marten, Fisher, Lynx and wolverine: survey methods

for their detection. Forest Service. United States Department of Agriculture. 163 pp.

4. Gese, E.M. 2003. Monitoring of terrestral carnivore populations (372-396). In: Gittleman, Funk

McDonald and Wayne (Eds.). Carnivore conservation. Cambridge. 675 pp.5. Vaughan, N., Lucas, E., Harris, S. & White, P.C.L. 2003. Habitat associatons of European hares (Lepus

europaeus) in England and Wales: implications for farmland management. J. Appl. Ecol. 40: 163-175.

DOI: 10.1046/j.1365-2664.2003.00784.x

6. Berg, R.L., Mc Donald, L.L. & Strickland, M.D. 1983. Distribution of mountain lions in Wyoming as

determined by mail questionnaire. Wildl. Soc. Bull. 11: 265-268.

7. Groves, C.R. 1988. Distribution of the wolverine in Idaho as determined by mail questionnaire.

Northwest Sci. 62 (4): 181-185.

8. Jamrozy, G. 1990. The occurence of the lynx in the Carpathian Mountains (south-eastern Poland)

according to questionnarie data. Acta Theriol. 35 (1-2): 162-164.

9. Rodrguez, A. & Delibes, M. 1990. El lince ibrico (Lynx pardina) en Espaa. Distribucin y

problemas de conservacin. [The Iberian Lynx (Lynx pardina) in Spain. Distribution and conservation

problems]. Coleccin Tecnica, ICONA. Madrid.

10. Rodrguez, A. & Delibes, M. 1992. Current range and status of the Iberian lynx Felis pardina

Temminck, 1824 in Spain. Biol. Conserv. 61:189-196.

DOI: 10.1016/0006-3207(92)91115-9
http://dx.doi.org/10.1046/j.1365-2664.2003.00784.xhttp://dx.doi.org/10.1046/j.1365-2664.2003.00784.xhttp://dx.doi.org/10.1016/0006-3207(92)91115-9http://dx.doi.org/10.1016/0006-3207(92)91115-9http://dx.doi.org/10.1046/j.1365-2664.2003.00784.x


9/10

31

11. SECEM 2001. Distribucin y estatus del turn (Mustela putorius) en Espaa: un anlisis basado

en encuestas. [Distribution and status of Polecat (Mustela putorius) in Spain: an analysis based on

hillsides]. Galemys13 (1): 39-61.

12. Carrier, J.A. & Beebee, T.J.C. 2003. Recent, substantial, and unexplained declines of the commontoadBufo bufo in lowland England. Biol. Conserv. 111: 395-399.

DOI: 10.1016/S0006-3207(02)00308-7

13. Gray, B.T. & Kaminski, R.M. 1993. Assessing a mail survey to estimate illegal waterfowl hunting.

Wildl. Soc. Bull. 21: 188-193.

14. Mazurkiewick, S.M., Boyle, K.J., Teisl, M.F., Morris, K.I. & Clark, A.L. 1996. Recall bias and

reliability of survey data: moose hunting in Maine. Wildl. Soc. Bull. 24(1): 140-148.

15. Crte, M. & Messier, F. 1987.Evaluation of Indices of Gray Wolf (Canis lupus) Density in Hard

Wood- Conifer Forests of Southwestern Quebec. Can. Field Nat. 101 (2): 147-152

16. Llaneza, L., Palacios, V., Uzal, A., Ordiz, A., Sazatornil, V., lvares, F. & Sierra, P. 2005. Distribucin

y aspectos poblacionales del Lobo Ibrico (Canis lupus signatus) en las provincias de A Corua y

Pontevedra. [Distribution and populational aspects of The Iberian Wolf (Canis lupus signatus) in the

A Corun and Pontevedra provinces]. Galemys 17 (N.E.): 61-80.

17. Llaneza, L., lvares, F., Ordiz, A., Sierra, P. & Uzal, A. 2004 Distribucin y aspectos poblacionales

del Lobo Ibrico en la provincia de Ourense. [Distribution and populational aspects of The Iberian

Wolf in the Ourense province]. Ecologa 18: 227-238.

18. Llaneza, L. & Ordiz, A. 2003. Distribucin y aspectos poblacionales del Lobo Ibrico (Canis lupus

signatus) en la provincia de Lugo. [Distribution and populational aspects of The Iberian Wolf (Canis

lupus signatus) in the Lugo province]. Galemys,15 (NE):55-66.

19. Cagnolaro, L., Rosso, D., Spagnesi, M. & Venturi, B.M. 1974. Inchesta sulla distribuzione del lupo(Canis lupus L.) in Italia e nel Canton Ticino e Grigioni (Svizzera). [Investigation into the distribution

of the wolf (Canis Lupus L.) in Italy in the Canton Ticino e Grigioni (Svizzera)]. Ric. Biol. Selv.; Lab.

Zoo. Apl. Caccia Bologna. 59: 1-9.

20. Blanco, J.C., Cuesta, L. & Reig, S. 1990. El lobo en Espaa: Una visin global. (69-93). [The Wolf

in Spain: general overview]. In: Blanco, J.C., Cuesta, L. & Reig, S. (Eds). El lobo (Canis lupus) en

Espaa. [The Wolf (Canis lupus) in Spain]. Coleccin Tcnica. ICONA. Madrid. 130 pp.

21. Llaneza, L. & Blanco, J.C. 2005. Distribucin y aspectos poblacionales del Lobo Ibrico (Canis lupus

signatus) en Castilla y Len. [Distribution and populational aspects of The Iberian Wolf in Castilla y

Len]. Galemys17 (N.E): 15-28.

22. Valverde, J.A. 1971. El lobo espaol. [Spanish Wolf]. Montes 159: 229-241.

23. Nez-Quirs, P., Garca, R. & Llaneza, L. 2007. Anlisis de la distribucin histrica del lobo (Canis

lupus) en Galicia: 1850, 1960 y 2003. [Analysis of Wolfs (Canis lupus) historical distribution in

Galicia]. Ecologa, 21: 195-205.

24. Filion, F.L. 1978. Increasing the effectiveness of mail surveys. Wildl. Soc. Bull.6: 135-141.

25. Llaneza, L., Rico, M., & Iglesias, J. 1998. Descripcin y resultados de varios mtodos de muestreo

para la deteccin y censo de Lobo Ibrico (Canis lupus signatus) en una zona de montaa. [Description

and results of various sampling methods for the detection and sensus of Iberian Wolf (Canis lupus

signatus) at a montainous site]. Galemys 10 (NE):135-149.

26. Hosmer, D.W. & Lemeshow, S. 2000. Applied logistic regression, 2nd ed. John Wiley & Sons, NewYork. 375 pp.

DOI: 10.1002/0471722146

27. Ministry of Environment, Lands & Parks 1998.Inventory Methods for Wolf and Cougar. Standards

for Components of British Columbias Biodiversity No. 34.

http://srmwww.gov.bc.ca/risc/pubs/tebiodiv/wolfsandcougars/Cited 25 March 2005.
http://dx.doi.org/10.1016/S0006-3207(02)00308-7http://dx.doi.org/10.1002/0471722146http://srmwww.gov.bc.ca/risc/pubs/tebiodiv/wolfsandcougars/http://srmwww.gov.bc.ca/risc/pubs/tebiodiv/wolfsandcougars/http://srmwww.gov.bc.ca/risc/pubs/tebiodiv/wolfsandcougars/http://dx.doi.org/10.1002/0471722146http://dx.doi.org/10.1016/S0006-3207(02)00308-7


10/10

32

28. Cea, A. 2000. Estudio del lobo (Canis lupus) en La Rioja mediante entrevistas (aos 1983-1994).

[Study of Wolf (Canis lupus) in La Rioja using interviews (1983-1994)]. Galemys 12 (NE): 201-208.

29. Gros, P.M. 1998. Status of the cheetah Acinonyx jubatus in Kenya: a Field interview assessment. Biol.

Conserv. 85: 137-149 DOI: 10.1016/S0006-3207(97)00135-3

30. Linnel, J.D.C., Swenson, J.E., Landa, A. & Kvam, T. 1998. Methods of monitoring European large

carnivores. A woldwide review of revelant experience. NINA Oppdragsmelding, 549: 1-38.

31. Filion F.L. 1980. Human surveys in wildlife management. In: Wildlife Management Techniques

Manual. Ed. S.D. Schemnitz. The Wildlife Society, Washington D.C., pp. 441-453

32. Gros P. M., Marcella J. Kelly & Caro, T.M. 1996. Estimating carnivore densities for conservation

purposes: indirect methods compared to baseline demographic data. Oikos 77:197-206.

33. Petrucci-Fonseca, F. 1990. O lobo (Canis lupus signatus Cabrera, 1907) em Portugal. Problemtica da

sua conservao. [The Wolf (Canis lupus signatus Cabrera, 1907) in Portugal. Conservation issues].

Ph D Thesis, Lisboa University 392 pp.

34. Lariviere, S., Jolicoeur, H. & Crete, M. 2000. Status and conservation of the gray wolf (Canis lupus)

in wildlife reserves of Quebec. Biol. Conserv. 94: 143-151.

DOI: 10.1016/S0006-3207(99)00185-8
http://dx.doi.org/10.1016/S0006-3207(97)00135-3http://dx.doi.org/10.1016/S0006-3207(99)00185-8http://dx.doi.org/10.1016/S0006-3207(99)00185-8http://dx.doi.org/10.1016/S0006-3207(97)00135-3

distribution of the iberian wolf in galicia. concordance between field sampling and questionnaires....

Documents