A c ta T h e r io lo g ic a 42 (2): 1 6 9 -1 7 7 ,1 9 9 7 .

PL ISSN 0 )0 1 -7 0 5 1

Habitat selection of genet Genetta genetta

in the mountains of central Spain

Emilio VIRGOS and Jorge G. CASANOVAS

Virgos E. and C asanovas J. G. 1997. H abitat selection o f genet G enetta gen etta in the
m ountains o f cen tral Spain. A cta Theriologica 42: 1 6 9 -1 7 7 .

G e n e t G en etta gen etta L in n aeu s, 1758 h a b ita t s election w as stu died th rou gh
m ;dden locations in a m ountain area o f central Spain both at m acrohabitat (habitat
types, land use, roughness, distance to villages, etc) and m icrohabitat scale (tree,
scru b, rock and p a stu re covers). A m on g the a v a ila b le h ab ita ts, gen et p referred
dehesas, due to their good trophic resources and shelter availability. G enet was scarce
over 1200 m etres, p roba b ly because o f th erm ic restriction s. The m ost im p ortan t
m icrohabitat variables selected as faecal sites w ere rock-cover (refuges) and shrub-
-cover (feeding places and refuges). O ther param eters o f functional significance related
to m id d e n lo ca tio n w ere s o cia l com m u n ica tion and te rrito ria lity . T h ere w as no
selection both sites w ith a high degree o f tree cover (m icroh abitat) and for sites near
to w atercourses.

K ey w ord s: G enetta genetta, habitat selection, m idden location, th erm ic restriction,

Spain

D epartm ent o f Anim al Biology I (Vertebrate Zoology), Faculty o f Biology, U niversidad

C o m p lu te n se , E -2 8 0 4 0 M a drid, S p ain , e-m a il: e v irg o s@ e u cm a x .sim .u cm .e s (E V );
C / B ernardina G arcía, 27, 28047 M adrid, Spain (JGC)

Introduction

Although the genet Genetta genetta Linnaeus, 1758 is relatively abundant in

most of its distribution area (Livet and Roeder 1987) the biology and ecology of
this species is still largely unknown. Most of the studies published focus on the
diet of this species (eg Calvino et al. 1984, Hamdine et al. 1993, Palomares and
Delibes 1991, Virgos et al. 1996) although other works focused on different
ecological aspects (Delibes 1974, Cugnasse and Riols 1984, Livet and Roeder 1987).
Microhabitat selection of faecal sites and the spatio-temporal ecology of this species
has recently been studied in Donana, SW Spain (Palomares 1993, Palomares and
Delibes 1994).
Habitat selection is considered to be a key aspect of animal activity, affecting
subsequent behavioral decisions and, ultimately, fitness (Orians and Wittenberger
1991). However, it is necessary to know the basic aspects of habitat choice before
testing their effects on population dynamics, breeding success and interspecific
interactions (Pulliam and Danielson 1991). In this context, habitat selection has

[ 169 ]
170 E. Virgos and J. G. Casanova

been viewed as a hierarchical process from the small spatial scale of feeding
patches to regional scales dealing with macroecological factors (Johnson 1980,
Morris 1987). Mediterranean mountains are highly suitable for these kinds of
studies due to their high environmental heterogeneity and the steep gradient of
habitats they present (Rivas-Martinez et al. 1987). Heterogeneous conditions and
environmental diversity in these areas offer an interesting opportunity to study
the different responses of individuals and populations to different but nearby
environments. On the other hand, it is necessary to attain baseline data in order
to be able to design scientific management programs for areas inhabited by genets.
This work describes the habitat selection patterns of genets at microhabitat
(faecal sites) and macrohabitat spatial scales from the Mediterranean mountains
of central Spain.

Material and methods

Stu d y area

The study area w as located in the Alto M anzanares Regional Park (41°40’N, 4°10’E) and its
surroundings, on the southern slope o f the G uadarram a m ountains (central Spain). The total area
studied covers 65,000 ha. All vegetation stages o f the central Spain m ountains w ere represented in
the study area. The low er level ranges between 6 0 0 -1 0 0 0 m a.s.l. and is covered by a m esom editer-
ranean holm oak Quercus ilex forest and dehesas, a typycal habitat w here the hum an-use (cattle)
creates a m osaic o f w oods and pastures. In holm oak forests the m ost im portant land-use is game.
The next stage (1 0 0 0 -1 7 0 0 m a.s.l.) was originally occupied by a Pyrenean oak Quercus pyrenaica
forest w hich has now been replaced by a m ixture o f different pine species. The level from 1 7 0 0 -2 0 0 0
m a.s.l. is covered by an orom editerranean S cot’s pine Pin us sylvestris forest and a scrubland
dom inated by Cytisus orom editerraneus. The two princpals land-uses w ere forestry and recreative
activities. Finally, the upper stage (2 0 0 0 -2 2 0 0 m a.s.l.) is a criorom editerranean pasture, rarely used
by genet and, therefore, not studied. Further inform ation about clim atic and vegetation types in the
study area can be consulted in Rivas-M artinez et al. (1987) and Font (1983).

S u rvey p ro ce d u res

Field w ork was carried out between 1989 and 1991, from October to April. The survey consisted
o f an exhaustive search o f the m iddens (scats am ounts) located in predictable places in the field
(R oed er 1978, 1980). Livet and R oeder (1987) use a sim ilar procedu re to stu dy genet h abitat
requirem ents in France and this procedure is also recom m ended for carnivore-habitat studies by other
authors (K endall et al. 1992, Clevenger 1993).
Field w ork was carried out by four people with experience in midden location. The w hole study
area w as divided into 5 x 5 km cells (n = 29), each classified follow ing habitat stratification criteria
(see Table 1 for habitat type). In order to m inim ize the error due to pseudoreplication, only the
m iddens located sufficiently far from each other (about 1000 m) were considered. In order to avoid
biases associated w ith scat data (Andelt and A ndelt 1984, Kruuk et al. 1986, Clevenger 1993),
selection patterns w ere studied w ith presence/absence data, although the m arking behaviour o f this
species (Roeder 1980) permits more accurate estim ations.
Variables 1 -7 (see Table 1) w ere quantified on maps and could be considered as descriptors o f
m a croh a b ita t environm ent: altitude (A LTIT ), an index o f roughness (RO U G ), distance to near
w atercourse (W ATER), distance to near village (VILL), distance to near road (ROAD), habitat-type
 Genet habitat selection 171

Table 1. D escription o f the m acrohabitat (1 -7 ) and m icrohabitat variables (8 -1 1 ) used in this study.

Variable Measure

1. ALTIT Height above sea level in each point

2. ROU G Num ber o f level curves intercepted in 500 m radius around each point
3. W ATER Minimun distance (in m) from each point to the nearest w atercourse
4. VILL Minimun distance (in m) from each point to the nearest village
5. ROAD Minimun distance (in m) from each point to the nearest road
6. H AB ITA T Habitat type: holm oak forest, dehesa - a typical landscape w ith woods and
pastures, and pine forest in each point
7. LAN D USE Type o f land use: cattle, forestry, recreative, and gam e in each point
8. TREE Tree canopy cover in 25 m radius around each point
9. SHRUB Shrub cover in 25 m radius around each point
10. ROCK Rock cover in 25 m radius around each point
11. HERB Herb cover in 25 m radius around each point

(H ABITAT), and land-use (LAND USE). The rem aining variables w ere m easured in the field and
concerned m icrohabitat features associated with defecation sites: tree cover (TREE), shrub cover
(S H R U B ), rock cover (RO C K ), and herb cover (H E R B ) (T able 1). S licin g o f m icroh a b ita t and
m acrohabitat variables has been considered as a useful tool to describe density-dependent habitat
selection patterns (M orris 1987). Variables belonging to the first set (m acrohabitat) w ere m easured
by m apping the m idden locations on a 1:50,000 topographical map. The second group o f variables
(m icrohabitat) w ere m easured in the field by visual estim ation w ithin circular-plots o f 25 m radius
around the m iddens. Availability o f the first set o f variables was quantified using 100 random points
scattered h om ogeneously over the topographical maps follow ing a stratified sam pling procedure. The
second group (m icrohabitat) w as quantified by carrying out 546 transects 500 m long follow ing a
stratified protocol too. The stratification procedure was made according to habitat availability in the
study area (60% holm -oak forest, 20% pine forests, 10% dehesas). In these transects, valu es o f
variables w ere visu ally estim ated w ithin circular-plots o f 25 m radius spaced at 125 m intervals.

D ata a n a ly se s

The existence o f selection in each variable was tested using a chi-square goodness o f fit test (Sokal
and R oh lf 1981). E xpected values were calculated from the availability data.
H ow ever, the m ultidim ensional characteristic o f a species’ en vironm ental requirem ents requires
a m ultivariate statistical approach, in order to avoid the problem o f non-independence o f variables.
Am ong all the new and old methods to analyze m ultivariate data, principal com ponent analysis (PCA)
can be used to stu dy the distribution of genet into an en vironm ental gradient generate previously by
availability data (Jam es and M cCullogh 1990). This procedure creates independent factors (ortho-
gonals) w hich are com posed by a linear com bination o f the original variables. Thus, this approach
allow s us to deal w ith correlated variables.
W e perform ed tw o separate analyses, the first w ith topographical and habitat-land use variables
(m acroh abitat variables) and, the second with m icrohabitat features associated with faecal sites
(m icrohabitat variables). This procedure was chosen for our study because o f the different ecological
significance and scale-m easure o f both variable types.
P rio r to th ese a n a ly se s va ria b les w ere tran sform ed as d ecim a l log a rith m ic (altitu d e and
distances), square root (roughness) and arcsine (covers) follow ing Zar (1984).
172 E. Virgos and J. G. Casanova

Results

U n iv a r ia te c h i-sq u a r e a n a ly sis

ALTIT, HABITAT and LAND USE were the only significant macrohabitat
variables. Genets prefers the 1 0 0 0-1 20 0 m a.s.l. interval. The dehesas was the

Table 2. Chi-square values for each variable, num ber o f genet m iddens (n ) and expected values (after
its environm ental availability) in each category.

Variable Expected Variable Expected

ALTIT LAN D USE

6 0 0 -8 0 0 1 5.2 Cattle 10 4.16
8 0 0 -1 0 0 0 12 7.8 Forestry 0 2.6
1 0 0 0-1 2 0 0 9 4.9 Recreative 5 6.93
> 1200 4 8.06 Game 11 11.7
%23 = 10.4, p < 0.05 X*3 = 8 .9 3,/> < 0.001

ROUG TREE
1 -2 10 10.1 0 -2 5 14 15.91
3 -4 5 5.46 2 5 -5 0 7 6.22
5 -6 7 3.12 > 50 5 3.86
>7 4 7.54 X22 = 0.64, ns
X23 = 6.61, ns

W A TER SHRUB
0 -2 00 10 10.66 0 -2 5 13.6
2 0 0 -4 0 0 8 7.28 2 5 -5 0 4.91
> 400 8 8.32 > 50 15 7.47
X22 = 0.25, ns X 2 = 17.79, p < 0.001

VILL ROCK
0 -1 0 0 0 4 6.24 0 -2 5 2 18.47
1 0 0 0 -2 0 0 0 5 6.5 2 5 -7 5 15 5.82
2 0 0 0 -3 0 0 0 8 6.5 > 75 9 1.71
> 3000 9 7.02 X^2 = 60.22, p < 0.001
%23 = 2.05, ns

ROAD HERB
0 -6 0 0 2 4.68 0 -2 5 23 19.27
6 0 0 -1 2 0 0 10 5.46 > 25 3 6.73
1 2 0 0 -2 4 0 0 4 5.91 X2 i = 2.78, ns
> 2400 10 9.88
X23 = 6.17, ns

H ABITAT
H olm -oak forest 13 15.86
D ehesa 10 2.34
Pine forest 3 4.68
X 22 = 22.77, p < 0.001
 G enet habitat selection 173

preferred HABITAT category and the cattle-use was the only significant LAND
USE one.
Among the microhabitat variables, we found significant selection for SHRUB
and R OCK ones. Genets preferred intermediate-high covers of both variables (see
Table 2 for details and statistics).

P C A from m a c r o h a b ita t d e sc rip to rs

Three factors were obtained, which explains 76.77% of the total variance
(Table 3). These factors were VARIM AX rotated to improve the biological inter­
pretation of the originals one. The first factor generates a gradient from elevated
and roughly regions of the study area (positive scores) to low-lying areas (negative
scores). The second, describes the distance to villages (VILL), with the distant

T able 3. R esu lts o f PCA analyses (VARIM AX rotation) from m acrohabitat (A) and m icrohabitat
variables (B)) used in this study. EIG EN V - Eigenvalues obtained from each PCA factor.

A B

PCA factor PCA factor

1 2 3 1 2 3

ALTIT 0.925516 0.190967 0.062309 TREE 0.046714 -0 .9 7 4 0 6 6 0.036040

R OU G 0.919506 0.070343 -0.006491 SHRUB -0 .9 0 1 1 2 7 0.207285 0.054157
W ATER 0.035157 -0.0 4 5 1 5 8 0.995316 ROCK -0 .0 6 3 4 8 4 0.045036 -0 .9 7 3 1 7 5
V ILL 0.161579 0.915981 -0.0 5 0 4 6 3 HERB 0.746603 0.271697 0.396818
ROAD 0.203975 0.420326 -0.1 0 6 7 7 4 EIGEN V 1.375659 1.067619 1.108766
EIG E N V 1.771020 1.059151 1.008527
%Total 34.3915 26.6905 27.7191
% Total 35.4204 21.1830 20.1705

points associated to positive scores. Finally, the last factor creates a gradient from
points near to watercourses (WATER) (negative scores) to more distant points
(positive scores).
Genets were associated with negative scores of the first factor (low-lying areas)
and thrid factor (WATER). When the second (proximity to villages) and third factor
were plotted, is possible observe an assocation of genet middens with areas near
to villages;. However, this relationship dissapears when it is compared to first
factor and, therefore, it is not plotted (Fig. 1).

P C A fro m m icro h a b ita t d e sc r ip to r s o f fa e c a l sites

We also obtained three VARIMAX rotated factors, which accounts 88.8% of

total variance (Table 3). The first factor describes a gradient from the pasture and
174 E. Virgos and J. G. Casanova

Fig 1. M idden distribution on the environm en­ Fig. 2. M idden distribution on the en vironm en ­
tal gradient generates from the PCA analysis tal gradient generates by PC A analysis from
o f m acrohabitat variables. Points mean indi­ m icrohabitat variables.
vidual genets.

open areas (positive scores) to shrub and closed ones (negative scores). The second
factor is related to a gradient of tree cover (with high covers associated to positive
scores). The last factor describes areas with different rock covers (high covers
associated to positive scores).
Genets were homogeneously distributed on the second factor (tree cover
gradient). Furthermore, the middens were concentrated on the negative scores of
the first factor (high shrub covers) and the positive scores of the third one (high
rock covers) (Fig 2).
 Genet habitat selection 175

Discussion

The first studies on the general biology of the genet (Delibes 1974, Cugnasse
and Riols 1984) revealed the flexibility of this species. In our study area, genets
also seem to follow this pattern, both with respect to diet (Virgos et al. 1996) and
the basic aspects of its habitat selection (this study). For all the variables studied
we found that the genet has a strong preference to living in basal habitats and
rarely above 1200 m a.s.l., in accordance with previous studies (Cugnasse and
Riols 1984, Livet and Roeder 1987, Ruiz-Olmo 1990). Most middens in these
low-lying areas were in dehesas (associated to cattle-use), whereas a few were
found in holm-oak forest. The dehesas are optimal habitats with abundance in
trophic resources supply (especially small mammals and birds, but also dung
beetles, E. Virgos and J. G. Casanovas, unpubl.) and shelter (hollow trees and
rocky places). Also, this choice reveals the affinity of this species for thermic places
as pointed out in earlier studies (Schauenberg 1966, Delibes 1974, Ruiz-Olmo 1990,
but see Desmet and Hamdine 1988). Despite the climatic wildness and richness
in trophic resources and shelter (rocky places) of areas between 6 0 0 -8 0 0 m
(holm-oak forests), these were not selected. This fact may be caused by carnivore
control techniques, and this is supported by the low richness and density of other
carnivore species (Casanovas and Virgos 1993). On the other hand, the genet
preferred higher areas than in France (Livet and Roeder 1987), where it lived in
altitudes between 0 -6 0 0 m a.s.l. and was rarely found above 800 m. A possible
explanation is the climatic differences between both areas, although, due to the
absence of availability resource information in the French data, we do not know
the selection criteria.
The PCA analysis reveals a tendency of genets to inhabit areas near to villages
(when we compare the second and thid factor in macrohabitat analysis), however,
this fact is explained by the association of this species to low-lying areas (due to
above suggestions), which were more man-populated (see the plot between the
first and second factor). Livet and Roeder (1987), pointed out a tendency to live
in areas with low human presence, although Delibes (1974) and Ruiz-Olmo (1990)
suggestions were in accordance with our data. On the other hand, the significance
of distance to watercourses is controversial. Chi-square approach reveals a lack
of selection for this variable. However, the PCA shows an association of middens
to areas near to watercouses. This factor were considered very important for some
researches (Cugnasse and Riols 1984, Livet and Roeder 1987), but this association
is not found elsewhere (Delibes 1974). Selection of areas near to watercourses may
enhance the availability and diversity of trophic resources (Livet and Roeder 1987).
The not selection of places with dense tree cover is remarkable, but has also
been recorded elsewhere (see Ruiz-Olmo 1990, Palomares and Delibes 1994). One
possible explanation is that the potential shelter role previously fulfilled by trees
is realized in the study area by rocks, which are very abundant both in the dehesa
and holm-oak forest. Rocks were a perfect shelter for genets in providing a warmth
176 E. Virgos and J. G. Casanova

(see Waechter 1975 for the stone marten) and undisturbed place. Moreover, the
genet usually selects rocky places well protected by shrubs, which have not only
a protective function but are also the habitat for some outstanding prey such as
wood mice Apodemus sylvaticus (Alcántara 1992, Virgos et al. 1996). On the other
hand, microhabitat features associated with middens could have a different
functional significance. Likely, middens have a social and territorial function
(Roeder 1978, 1980 for genets, Macdonald 1980 for a review from carnivore
communities) and, therefore, their location could be aimed at maximizing detection
by conspecifics. The high preference for tall structures which are available both
here and elsewhere (trees, high rocks, etc) is due to the same reason. On the other
hand, the two alternatives may be not exclusives and, therefore, a two-significance
role (social communication-shelter) is possible to be associated to midden places.
This subject requires future behavioral research.
In summary, genets selected areas lower than average probably due to thermic
restrictions, although the holm-oak forests could be not selected due to human
persecution. Among the microhabitat variables, places with high rock and scrub
covers were selected.
A cknow ledgem ents: W e wish to thank the follow ing people for their help and contributions to this
work: T. Blázquez, F. J. Sam blás, D. García, Y. Cortés, M. A. García, L. Suárez and R. M artinez for
field advices; G. P. Farinós, G. G. Nicola, M. M orales and V. Virgos for translation; F. Palom ares for
som e references and J. L. Telleria for helpful suggestions on early drafts on the m anuscript. Three
anonym ous referees gave valuable com m ents on the first version o f the m anuscript. This w ork was a
contribution to PB 92/0238 D G ICYT project.

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R eceived 17 July 1995, revised 10 February 1997, accepted 1 M arch 1997.