Introduction

Habitat models are useful tools for helpingmanagers and researchers to predict the distri-bution or abundance of a species (Litvaitis et al.

1994, Morrison et al. 1998, Guisan and Zimmer-mann 2000, Rodríguez et al. 2007).

We studied the distribution and abundance ofthe Iberian hare Lepus granatensis Rosenhaeur,1856 and the European wild rabbit Oryctolagus

cuniculus (Linnaeus, 1758). These species are ofkey ecological importance in Iberian ecosystems,mainly in the Mediterranean area (Delibes andHiraldo 1981, Rogers 1981, Serrano 2000, Angulo

[73]

Acta Theriologica 55 (1): 73–79, 2010.

PL ISSN 0001-7051 doi: 10.4098/j.at.0001-7051.018.2009

Modelling habitat use by Iberian hare Lepus granatensis and

European wild rabbit Oryctolagus cuniculus in a mountainous

area in northwestern Spain

Luis TAPIA, Jesús DOMÍNGUEZ and Luis RODRÍGUEZ

Tapia L., Domínguez J. and Rodríguez L. 2010. Modelling habitat use byIberian hare Lepus granatensis and European wild rabbit Oryctolagus cuniculusin a mountainous area in northwestern Spain. Acta Theriologica 55: 73–79.

We developed broad-scale habitat use models for the distribution of theIberian hare Lepus granatensis Rosenhaeur, 1856 and the relative abundanceof the European wild rabbit Oryctolagus cuniculus (Linnaeus, 1758) in a moun-tainous area in north-western Spain, the Baixa Limia Site of CommunityImportance (SCI). For the Iberian hare, the most parsimonious model includedaverage altitude, average slope, scrub-pasture area and length of border betweenscrublands and forests. Four variables were consequential, 2 of them with apositive sign for the presence of the hare (average altitude and scrublandarea) and 2 with a negative one (average slope and length of border betweenscrubland and forest). For the European wild rabbit the most parsimoniousGeneral Linear model obtained included average altitude and scrub-forestedge. Only 2 predictors were consequential, both with a negative sign on therelative abundance of wild rabbits: the average altitude and length of borderbetween scrubland and forest. The models obtained indicated the need of man-agement actions to favour open habitats with suitable vegetation structure,avoiding extensive reforestation. A revision of the current hunting plans inthe protected area of Baixa Limia is required.

Departamento de Zoología y Antropología Física, Facultad de Biología, Universidad de Santiagode Compostela, Campus sur, s/n 15782, Galicia, Spain, e-mail: luis.tapia@usc.es (LT, JD);Departamento de Edafología y Química Agrícola, Facultad de Biología, Universidad de Santiagode Compostela, Campus sur, s/n 15782, Galicia, Spain (LR)

Key words: Lepus granatensis, Oryctolagus cuniculus, game species, habitat use,Spain

2003, Calvete et al. 2004, Delibes-Mateos et al.2008). Moreover, they are important game spe-cies that require appropriate management inSouthern Europe, a region with a strong tradi-tion of hunting (Ballesteros 1998). Data on thehabitat selection by these species in Europeancountries were obtained mainly from studies atthe microhabitat level (Duarte 2000, Herranz et

al. 2000, Caruso and Siracusa 2001, Nadal 2001,Angulo 2003, Carvalho and Gomes 2003, Virgóset al. 2003, Monzón et al. 2004, Lombardi et al.

2007). There is little information available re-garding the north-western area, where naturalpopulations are generally depleted and of lowdensity (Carvalho and Gomes 2003, Monzón et

al. 2004), mainly due to poor habitat conditionsand disease (Fenner and Ross 1994, Villafuerteet al. 1995, 1998, Calvete et al. 2004).

The development of a habitat suitabilitymodel for the species under study is an impor-tant step in establishing management and re-covery plans (Rodríguez et al. 2007).

The aim of this study was to develop broad-scale habitat models for the Iberian hare and theEuropean wild rabbit in a mountain environ-ment in the north-western part of the Iberianpeninsula using presence/absence and relativeabundance indices, respectively, as dependentvariables, and habitat predictor variables. Thestudy will contribute to new insight into habitatselection patterns for these species at a land-scape scale (Buckland and Elston 1993, Rushtonet al. 2004).

Study area

The study area comprises the mountainous environ-ments above 800 m a.s.l. in the Baixa Limia Site of Commu-nity Importance (SCI) (34 627 ha), which includes BaixaLimia-Serra do Xurés Natural Park (20 920 ha). The areaextends along the south-western mountains in the provinceof Ourense (Galicia, NW Spain; 42°00’N–8�00’W) and bor-ders the National Park of Peneda-G�rés in Portugal. TheSpanish and the Portuguese protected areas cover an areaof 106 627 ha altogether. The zone is a mountain range,with peaks of 1500 m, formed predominantly by graniterocks. Currently, the number of human population densityin the area is quite low, although the landscape has beenintensely affected by human activity. The climate in thearea is temperate oceanic sub-Mediterranean, with a meanannual temperature of 8–12 �C and a mean annual precipi-

tation of 1200–1600 mm, which involves a significant watershortage in summer (Martínez-Cortizas and Pérez-Alberti1999). The most common types of vegetation are the scrubcommunities (Ulex spp., Chamaespartium tridentatum,Erica spp., Genista spp. and Cytisus spp.). Woods are veryfragmented, and dominated by oaks (Quercus robur, Q.

pyrenaica) and pines (Pinus pinaster, P. sylvestris). Allplant communities in the study area are under strong pres-sure from frequent deliberate fires and extensive livestockgrazing (Pulgar 2003). This leads to a control of the scrubcover and the generation and maintenance of pasture areas.

Material and methods

For the Iberian hare, systematic ground surveys using a1 � 1 km grid were carried out, in spring and summer, from1997 to 2000, to detect its presence (droppings and directobservation). The information was completed using datafrom semi-quantitative surveys (Tapia and Domínguez2003), obtained with night-road surveys and night spotting(Gortázar et al. 2007). Observations provided by other biolo-gists working in the area and gamekeepers from the SCIBaixa Limia have also been used. It was assumed that thedistribution of the Iberian hare in the area is known withfull precision, with no false absences (Hirzel et al. 2002,Miller et al. 2004).

For the European wild rabbit, a semi-quantitative sur-vey in areas between 800 and 1500 m a.s.l. was carried outbecause most of the study area (70%) is situated above thisaltitude, and the habitat is more homogeneous. Relativeabundance indices were obtained for the spring-summer pe-riod, and were based on the number of faecal pellets found,assuming a direct relationship between this parameter andthe abundance of rabbits (Wood 1988, Palomares 2001).Due to the difficulty of surveying all the 1 � 1 km squaresthat were higher than the 800 m a.s.l. cut-off point (184grids), a prior random selection of 89 sites was made andthese squares were sampled in the spring and summer of1998 in the following way: two parallel routes of 600 mlength, separated by 300 m and oriented N–S, were made ineach of the selected squares. On each route, six transects of10 m length and 1 m width were made, along which thenumbers of all individual faecal pellets encountered werecounted. The transects were spaced 100 m apart. The start-ing point of each route was located 300 m from the ordinateaxis and 100 m from the abscissa of the lower left quadrantof each 1 � 1 km square selected. The exact location of thispoint was identified using a GPS. The number of transectsfor each 1 � 1 km square was 12, so that the effective sur-face surveyed was 120 m2. The abundance index obtainedcorresponded to the number of pellets/10 m2 per 1 � 1 kmsampled square.

For the habitat selection analysis, a 2 � 2 km grid wasused, including the information obtained in the 1 � 1 kmsquares sampled and allowing an analysis at the macro-habitat level (Ruiz-Martínez 1996, Morrison et al. 1998,Nadal 2001). The 2 � 2 km grid was chosen because theproportion of 1 � 1 km grid squares in which Iberian harewas present was low, and also because the cartographyused lost resolution at small scales. In order to avoid

74 L. Tapia et al.

problems derived from incomplete sampling, those 2 � 2 kmsquares of which less than 50% of the surface area wasincluded within the boundaries of the Baixa Limia SCI ormore than 50% was included in Portugal, were ruled out.The final sample size for the habitat models (2 � 2 kmsquares) was n = 93.

Geographic Information System (GIS) analysis

The model for the European wild rabbit was developedusing the index of abundance of pellets in a 1 � 1 km gridat an altitude higher than 800 m a.s.l. The average value ofabundance in the 2 � 2 km grid in the Baixa-Limia SCI wascalculated from the pellet abundance in the 2 � 2 km grid,using the Spatial Analyst Extension in Geographic Informa-tion System GIS-ArcView 3.1. (Environmental Systems Re-search Institute, Inc., Redlands, CA). This was finally thedependent variable used. This model was based on spatialextrapolation and interpolation techniques (Miller et al.

2004), in which the values of the dependent variable wereestimated on the basis of the geographical location with re-gard to sampled neighbouring points (Maurer 1994, Seoaneand Bustamante 2001). The abundance of rabbits was esti-mated using the “Inverse Distance Weighted” approach.This interpolation estimates the value for a point from theaverage value of the values of the sampled points in its vi-cinity. We decided to use a linear interpolator, assumingspatial dependence among points close to one another.There are two main linear interpolators: Inverse DistanceWeighted (IDW) and Spline interpolators. Splineinterpolator is better for gently varying surfaces and it isnot appropriate if there are large changes in the surfacewithin a short horizontal distance since it trends to overes-timate the values of the predicted variables. We thereforedecided that IDW was the best function for estimating theaverage value of abundance of rabbits. The IDW methoduses both the values of the points in the neighbourhood andthe values of points contained at a fixed radius to estimatethe value of a certain unsampled point. This approach mayfail to find neighbours if large spatial gaps exist in the data,so it will be unable to interpolate the whole surface appro-priately. We used a default value of 12 neighbouring pointsto ensure that a large number of samples were influencingon the final estimated values, with the highest effect onunsampled points being that of the nearest sampled point.We did not calculate the sensitivity of neighbourhood size inthe results, but we assume that the use of a lower numberof neighbours may lead to errors due to possible anisotropyin the distribution of data. Due to the limitations of theavailable digital cartography resolution, 6 predictors wereselected. They represented different land use (scrub-pas-ture area and forest area), degree of human/anthropogenicinfluence (area of settlements), topographic irregularity(mean altitude and mean slope) and habitat heterogeneity(scrub-forest edge), and values for each variable were deter-mined for each 2 � 2 km plot studied. These independentvariables were selected taking into account the habitat re-quirements of the two lagomorph species (Ballesteros 1998).The information related to the different environmentalvariables was taken from 1:50 000 digital maps with a reso-lution of 250 � 250 m using GIS-ArcView 3.1.

Continuous variables (ie, mean slope and mean altitude)were estimated from a digital elevation model at 250 m reso-lution. The remaining variables were obtained directly by aGIS using vectorial data. Scrubland and pasture surfaceswere determined using data from the CORINE Land CoverProject (EEA 1994). Scrubland and pasture surfaces are oftenintermixed so they were treated as one type of cover becausethey could not be distinguished at the spatial resolution used.All forest ecosystems were also treated as the same type, in-dependently of their tree species composition. Forests andscrubland-pastures represented almost 90% of the total studyarea. Forest information was obtained from the Third ForestSpanish National Inventory (D.G.C.N. 2000).

Statistical analysis

Generalized linear models (GLM) were used for bothspecies. In case of the Iberian Hare, a binary logistic modelwith binomial distribution and logit link was employed. Incase of the wild rabbit, a linear model with normal distribu-tion and identity link was used.

The best overall model is chosen from a group of well-de-fined candidate models (hypothesis) with biological meaning(Anderson et al. 2000, Seoane and Bustamante 2001, Johnsonand Omland 2004). For both species, all models of main ef-fects combining the 6 selected predictors were considered. Thebest model was selected using the corrected Akaike informa-tion criterion (AICc). This expression of AIC was used becausen/K < 40 (Anderson et al. 2000). For each model i the valuesof AICc and �i were obtained, where �i = AICi–AICminimum

model. Generally, models with �i values � 2 have strong sup-port, whereas those with values > 10 have little support. Theweight of each model was calculated as wi:

wi =exp

exp

��

��

�

�

��

��

�

�

�

12121

�

�

i

r

R

r

(Anderson et al. 2000)

The sum of all weights equals to the unit, and the valueof each wi indicates the model i is the best overall Kulback-Leibler model (Anderson et al. 2000, Johnson and Omland2004). Using the Akaike Information Criterion, no model isthe null model, nor is a significance level established a

priori. The importance of each predictor is obtained byadding the Akaike weights to the models in which thatvariable is present (Burnham and Anderson 1998). Theaddition of the weights of each variable was consideredconsequential when �wi > 0.5 (Taylor and Knight 2003).

All analyses were performed using the statistical pack-age SPSS v16.0.

Results

Presence of Iberian hares was detected in 16of the 2 � 2 km squares (17% of the 93 total

Lagomorphs habitat in northwestern Spain 75

squares). The most parsimonious model in-cluded average altitude, average slope, scrub-pasture area and length of border betweenscrublands and forests (Table 1). Another modelthat included all predictors except for the forestarea, was also competent (�i < 2). Four variableswere consequential, 2 of them with a positivesign of the presence of the species (average alti-tude and scrubland area) and 2 with a negativeone (average slope and length of border betweenscrubland and forest) (Table 2).

For the European wild rabbit, the faecal pel-let counts were positive in 31 of the 2 � 2 kmsquares (33.7% of the 93 total squares). Themost parsimonious model obtained included av-erage altitude and scrub-forest edge. Two othermodels were competent (�i < 2) (Table 1). Only 2predictors were consequential, both with a nega-tive sign on the relative abundance of wild rab-bits: the average altitude and length of borderbetween scrublands and forests (Table 2).

Discussion

Modelling wildlife-habitat relationships fromdistribution or abundance data by consideringthese as indicative of habitat quality, involvesthe assumption that species are selective intheir choice of habitat (Cody 1985) and that it ispossible to find correlations between environ-mental variables and distribution and abundance(Buckland and Elston 1993). Nevertheless, thismay not always be the case and may lead todifficulties in model development even in undis-turbed habitats (Van Horne 1983, Anderson andGutzwiller 1994, Beutel et al. 1999). In thepresent study we are aware that other variables,not considered, may be better predictors andprovide more reliable models. However, many ofthese variables (trophic availability, rabbit re-stocking, hunting pressure) were not consideredand would be difficult to measure at the scaleused, possibly even complicating the models.

76 L. Tapia et al.

Table 1. Summary of model selection results for presence of Iberian hares and abundanceof European wild rabbits in the 2 � 2 UTM grid in the Baixa-Limia SCI (n = 93). Modelswith �i < 2 are shown and ranked by descending Akaike weights (wi). Predictors: (1) areaof human settlements, (2) average altitude, (3) average slope, (4) area of scrubland andpastureland, (5) area of forests, (6) length of border between scrublands and forests.

Species MODEL AICc i wi

Iberian hare {2,3,4,6} 72.459 0 0.205{1,2,3,4,6} 74.261 1.802 0.083

European wild rabbit {2,6} 412.752 0 0.237{2,5,6} 414.229 1.477 0.113{2,3,6} 414.724 1.972 0.088

Table 2. Relative importance of effect magnitudes (calculated by adding for each variable the Akaikeweights for all models) of the predictive variables used for the modelling the Iberian hare presence/ab-sence and European wild rabbit abundance in the Baixa Limia SCI in the 2 � 2 UTM grid. In bold, val-ues over 0.5 considered as consequential. Predictor numbers as in Table 1.

SpeciesPredictors

1 2 3 4 5 6

Iberian hare –0.306 +0.846 –0.868 +0.676 –0.336 –0.691European wild rabbit –0.220 –0.999 +0.272 –0.310 –0.414 –0.839

In the case of the Iberian hare, there werelow values of abundance with associated conser-vation problems (Tapia and Domínguez 2003).The models obtained seemed to reflect the habi-tat requirements satisfactorily in terms of vege-tation structure at the scale employed. Thehares selected areas of high altitude, with gentleslopes, dominated by scrub-pasture areas. Thispattern of habitat use coincides with the re-quirements reported in mountainous areas inthe North of the Iberian peninsula (Castellsand Mayo 1993, Soriguer and Palacios 1996,Ballesteros 1998, Carro and Soriguer 2002). It isimportant to point out the abundant presence ofextensive livestock in most of mountainous areaswith presence of hares, which can have importantmanagement implications for the species.

The current status of wild rabbit in the studyarea may be the result of factors other than poorhabitat conditions, such as hunting manage-ment and disease (Tapia 2004). The relationshipbetween the abundance of rabbits and the habi-tat types may therefore be distorted (Soriguerand Palacios 1996, Vargas 2002, Angulo 2003,Carvalho and Gomes 2003). On the basis of theresults of the pellet counts, the environmentalvariables selected in the abundance modelsseem to indicate that the abundance of the wildrabbit may be favoured by the existence of areaswith scarce forest surface at a lower altitude.This habitat selection model may possibly corre-spond to a higher availability of open land withthe presence of appropriate pasture areas forfeeding and better conditions for the establish-ment of warrens (Ballesteros 1998, Purroy andVarela 2003, Lombardi et al. 2007). With the in-crease in altitude, the geology and topographywould imply a lower probability of detecting therabbits, as occurs in the rest of the Iberian pen-insula (Castells and Mayo 1993, Purroy andVarela 2003). It is also likely that other vari-ables which were not considered, such as vegeta-tion cover, edaphology (influence of soil type),microclimate and human activities (hunting,wildfires, habitat loss, restocking of huntingspecies), are involved in the patterns of habitatuse. All of these seem to determine the distribu-tion and abundance of the wild rabbit at finerscales of analysis (Angulo 2003, Carvalho and

Gomes 2003, Monclús and De Miguel 2003,Virgós et al. 2003, Lombardi et al. 2007). Thehabitat use models obtained in this study partlycoincide with those obtained for a neighbouringarea in Portugal (Carvalho and Gomes 2003,Monzón et al. 2004) and mosaic ecosystems inthe Centre and South of the Iberian peninsula(Herranz et al. 2000, Angulo 2003, Lombardi et

al. 2003, Virgós et al. 2003, Lombardi et al.

2007). However, the physiognomy, edaphic andphytosociological characteristics, as well as hu-man use, census methodology, modelling tech-niques and the scale of analysis used weredifferent from those in the present study. Thissuggests the necessity of combining fine andbroad habitat scales of analysis, to show in moredetail the ecological requirements of the wildrabbit in the Northwestern Iberian Peninsula.

The models obtained for both species indi-cated the need to favour open habitats with suit-able vegetation structure, avoiding extensivereforestation. A clearing of scrub (above 800 ma.s.l.) and the maintenance of extensive live-stock to favour the pasture are recommendedas management habitat actions for the hares(Thompson et al. 1995). The selection of thepatches to be cleared in the suitable habitats forhares should be made in accordance with therecommendations of a botanical expert, to avoidharming species of high interest in the area(Pulgar 2003). The ecotone interface betweenscrub and open herbaceous formations increasesthe density of wild rabbits, by providing a combi-nation of high trophic availability and protectivescrub cover (Villafuerte and Moreno 1997,Carvalho and Gomes 2003, Lombardi et al.

2003). In some areas of the Baixa-Limia SCI, anoverhunting of both species could currently oc-cur. A revision of the current hunting plans inthe protected area is required.

Acknowledgements: This study was financed by the Consel-lería de Medio Ambiente (Xunta de Galicia). We express ourgratitude to M. Romeu, X. Calvo, M. Vidal and I. Quinterofor their collaboration in the field work, to C. Rouco (Do�anaBiological Station, Spanish Research Council, CSIC) and V.Sel�s (Norwegian University of Life Sciences) for their com-ments and improvements of this manuscript. We thank F.Palacios (Spanish Research Council, CSIC), R. Soriguer andF. Carro (Estación Biológica de Do�ana, Spanish ResearchCouncil, CSIC) for helpful comments and improvements onthe lagomorphs census techniques.

Lagomorphs habitat in northwestern Spain 77

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Received 19 March 2009, accepted 10 November 2009.

Associate editor was Magdalena Niedzia³kowska.

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