233
Animal Biodi e si y and Conse a ion 37.2 (2014)
© 2014 Museu de Ciències Na u als de Ba celona
ISSN: 1578–665 X
eISSN: 2014–928 X
Della io e, C. M., Rouco, C., Muñóz Vallés, S. & Gallego Fe nández, J. B., 2014. Seasonal habi a use by
he Eu opean abbi (O yc olagus cuniculus) in a coas al dune sys em in SW Spain. Animal Biodi e si y and
Conse a ion, 37.2: 233–242.
Abs ac
Seasonal habi a use by he Eu opean abbi (O yc olagus cuniculus) in a coas al dune sys em in SW Spain.—
We s udied habi a use by he wild Eu opean abbi (O yc olagus cuniculus) in a coas al sand dune sys em in he
sou h–wes e n Ibe ian peninsula. Ou goals we e o de ine he use o his habi a by abbi s in ela ion o ood
and shel e a ailabili y be ween seasons. Rabbi densi y, ood a ailabili y and e uge abundance we e analysed
using mul iple eg ession analyses. We ound ha , independen ly o season, habi a selec ion was p incipally
ela ed o co e by he woody sh ub Re ama monospe ma which abbi s use bo h as a ood esou ce and as
p o ec ion agains p eda o s. Al hough i is an in asi e na i e plan , he bene i s ha R. monospe ma p o ides
o he wild abbi popula ion should be aken in o accoun when deciding wild abbi managemen s a egies.
Key wo ds: Re ama monospe ma, Ibe ian peninsula, Rabbi p eda ion isk, Feeding si e, Seed dispe sal
Resumen
Uso es acional del hábi a po pa e del conejo de mon e (O yc olagus cuniculus) en un sis ema de dunas
cos e as en el su oes e de España.— Es udiamos el uso del hábi a po pa e del conejo de mon e sil es e
(O yc olagus cuniculus) en un sis ema de dunas cos e as del su oes e de la península ibé ica. Nues os ob�
je i os ue on de ini la u ilización que los conejos hacen de es e hábi a en elación con la disponibilidad de
alimen os y cobijo en e es aciones. Se analiza on la densidad de conejos, la disponibilidad de alimen os y la
abundancia de e ugios median e análisis de eg esión múl iple. Obse amos que, independien emen e de la
es ación, la selección del hábi a es aba p incipalmen e elacionada con la cubie a del a bus o leñoso Re ama
monospe ma, que los conejos u ilizan como uen e de alimen o y como p o ección con a los p edado es. A
pesa de que se a a de una plan a na i a in aso a, los bene icios que R. monospe ma apo a a la población
de conejos sil es es debe ían ene se en cuen a a la ho a de decidi cuáles deben se las es a egias de
ges ión pa a es a especie.
Palab as cla e: Re ama monospe ma, Península ibé ica, Riesgo de p edación, Á ea de alimen o, Dispe sión
de semillas
Recei ed: 5 X 14; Condi ional accep ance:20 XI 14; Final accep ance: 1 XII 14
Claudia. M. Della io e, Uni e sidad Nacional de Río Cua o, Có doba, A gen ina.– Ca los Rouco, Landca e
Resea ch, P. O. Box 1930, Dunedin 9054, New Zealand.– Sa a Muñóz Vallés & Juan B. Gallego Fe nández,
Dep o. de Biología Vege al y Ecología, Uni . de Se illa, España (Spain).
Co esponding au ho : Ca los Rouco. E–mail: [email p o ec ed], [email p o ec ed]
Seasonal habi a use
by he Eu opean abbi
(O yc olagus cuniculus) in a coas al
dune sys em in SW Spain
C. M. Della io e, C. Rouco, S. Muñóz Vallés
& J. B. Gallego Fe nández
234 Della io e e al.
In oduc ion
Habi a selec ion is a i al decision o animals as i
a ec s ood, shel e and ep oduc ion (Ha gis & Mc�
Cullough, 1984; Pulliainen & Tunkka i, 1987; Mo ison
e al., 1998; Bond e al., 2002). This decision can be
in luenced by species’ li e his o y ai s, indi idual cha�
ac e is ics such as age and sex, and en i onmen al
ac o s such as compe i ion, p eda ion p essu e and
seasonal ood supply (e.g. Bos & Ca hew, 2003;
Lomba di e al., 2007; He nandez–Aguila e al., 2013;
Lisón e al., 2014).
O e he las decade, wildli e esea che s ha e
made g ea e o s o be e unde s and he p essu es
ac ing on p ey species o selec ce ain habi a ypes
and he implica ions o a oiding p eda o s. Se e al
s udies ha e shown he ecological lexibili y o he
Eu opean abbi (O yc olagus cuniculus) in Medi e �
anean ecosys ems, and i s wide plas ici y o adap i s
beha iou o p eda o p essu e and ood a ailabili y
(Jaksic & So igue , 1981; Lomba di e al., 2003, 2007;
S o , 2003; Sa men o e al., 2012).
The Eu opean abbi is a key p ey species in Sou h�
e n Eu ope (Delibes–Ma eos e al., 2008a) wi h g ea
capaci y o colonize di e en habi a ypes (Thompson
& King, 1994). On he Ibe ian Peninsula, he abbi has
a wide dis ibu ion and i has been obse ed in di e se
ecosys ems, om Medi e anean sc ublands o dehesa
ag oecosys ems, moun ainous a eas, and coas al sand
dunes (e.g. Rueda e al., 2008a, 2008b; Ba io e al.,
2009; Della io e e al., 2009; Tapia e al., 2010, 2014).
The coas al sand dune ecosys em ep esen s 40% o
he 7,880 km o he Ibe ian coas (Ley e al., 2011)
and abbi popula ions ha e been obse ed in almos
all such sand dune a eas (Villa ue e, 2002), bu hey
ha e been poo ly s udied o da e. The lack o in e es
could be because hese coas al sand dunes hold no
hun ing in e es and house no endange ed p eda o s.
In his s udy we ocused ou esea ch on he dune
sys em o 'La Flecha de El Rompido' in he sou hwes
o he Ibe ian Peninsula. Mos dune sys ems in his
a ea a e cu en ly expanding and La Flecha dune
sys em is g owing by mo e han 30 m pe yea a i s
eas e n edge (Gallego–Fe nández e al., 2006). I is
inhabi ed by a la ge popula ion o abbi s (Della io e,
2007). Ha es (Lepus g ana ensis) a e also p esen in
his ecosys em, bu a a lowe densi y, and ed ox
(Vulpes ulpes) is he main p eda o (Della io e, 2007).
P e ious s udies ha e es ablished ha abbi
habi a selec ion is based on wo main d i e s: ood
p o ision and p eda o a oidance (e.g. a p e e ence
o shel e ed habi a s) (Lomba di e al., 2007; Sa illi &
Bagliacca, 2010). The e o e, o de ine habi a selec�
ion by abbi s in a coas al sand dune sys em in he
sou hwes e n Ibe ian peninsula we we e pa icula ly
in e es ed in assessing he e ec o ood and shel e
a ailabili y on habi a use and also habi a use in di �
e en seasons. Ou s udy o habi a use in sand dune
a eas was designed o con ibu e o unde s anding
he abbi ’s huge capaci y o adap a ion o a wide
ange o en i onmen s and he lexibili y o i s spa ial
beha iou . Such in o ma ion could be o alue when
de e mining managemen s a egies o he species.
Ma e ial and me hods
S udy a ea and en i onmen al uni s
This s udy was conduc ed on El Rompido spi , on he
seawa d side o he Pied as Ri e es ua y (Huel a
P o ince, SW Spain, 37º 12' N, 7º 10' W). The spi
ex ends eas wa ds o 12 km, unning pa allel o he
coas , wi h a wid h anging be ween 300 and 700 m. The
a ea consis s o 527 ha o na u al sandy soil, o which
293 ha is composed o inne s abilized dunes ( ig. 1).
The clima e has a Medi e anean pa e n o we win e s
and d y summe s, wi h a mean annual empe a u e o
18.2ºC and mean annual ain all o 620 mm.
The s udy a ea has a ious ege a ion communi ies
ela ed o local geomo phology, including he beach
and ac i e dune sys em, sal ma shes, inne s abili�
zed dunes, we dep essions, and idal swales. The
ac i e dune zone is mainly co e ed by Ammophila
a ena ia (ma am g ass) and Elymus a c us (sand
couch–g ass). Due o ad e se condi ions, ege a ion
is spa se and low. The e is a ege a ion g adien in�
land acco ding o local en i onmen al condi ions. The
inne s abilized dune zone is he main spi su ace
a ea (56%) and is la gely co e ed by he woody sh ub
Re ama monospe ma (b idal eil b oom). Re ama mo-
nospe ma can g ow up o 3.5 m in heigh wi h a iable
co e . The plan occu s wi h chamaephy ic species
such as Thymus ca nosus, A emisia c i hmi olia, and
Helych ysum pica dii, which show di e se dis ibu ion
pa e ns along he spi . The R. monospe ma canopy
con ibu es o he high numbe o he baceous plan s,
la gely consis ing o win e annuals. We dep essions
a e co e ed by hyg ophy ic ege a ion (usually no
exceeding 1.5 m in heigh ), domina ed by he pe�
ennial Sci pus holoschoenus and/o Juncus acu us
and, occasionally, Sci pus ma i imus (depending on
wa e a ailabili y). These we dep essions a e also
colonized by se e al annual he baceous species,
especially win e annuals. Tidal swales a e dep es�
sions be ween dune idges connec ed wi h he i e
channel and domina ed by halophy ic ege a ion ( o
de ails see Gallego Fe nández e al., 2006; Annex).
We used geomo phologic ea u es and ege a ion
composi ion o classi y he s udy a ea in o 10 en i on�
men al uni s: (1) dune sc ubland, (2) R. monospe ma
wi h Thymus ca nosus, (3) R. monospe ma wi h A e-
misia c i hmi olia, (4) R. monospe ma wi h Helich ysum
pica dii, (5) inland dunes wi h R. monospe ma, (6) we
dep essions, (7) ac i e dune a ea, (8) idal swales, (9)
sal ma shes, and (10) sandy sho e ( able 1, also see
Gallego Fe nández e al., 2006, Della io e e al., 2008).
Sampling design and da a collec ion
Using a s a i ied andom sampling model, 146 poin s
we e selec ed h oughou he s udy a ea on he basis o
he en i onmen al uni s p esen . The numbe o poin s in
each en i onmen al uni was es ima ed acco ding o he
a ea o each uni and aking a minimum o 10 poin s in
he smalles uni (dis ance be ween poin s anged om
10–55 m). Poin s in each uni we e selec ed based on
a andomly gene a ed able o geog aphic coo dina es.
Animal Biodi e si y and Conse a ion 37.2 (2014) 235
Each poin was su eyed in win e (Janua y–Ma ch) and
sp ing (Ap il–June) o 2004, and du ing he summe
(July–Sep embe ) and au umn (Oc obe –Feb ua y) o
2005. We measu ed abbi ela i e abundance and ood
and e uge a ailabili y a each sampling poin pe season.
Rabbi ela i e abundance
Pelle coun s a e conside ed an e ec i e me hod o
es ima ing abbi ela i e abundance and o e alua �
ing habi a use by lagomo ph species (Mye s, 1955;
Jaksic e al., 1982). In consequence, abbi ela i e
abundance was assessed by pelle coun s in dung
clea ance plo s (Paloma es, 2001; Fe nández de Simon
e al., 2011a). One pe manen dung plo o 0.5 × 0.5 m
was placed a each o he 146 sampling poin s. E o s
we e made o a oid la ines and si es unde sh ubs
because hey can be used as e uge o es ing si es.
Each plo was cleaned 25–35 days be o e pelle col�
lec ion and coun s. To s anda dise all pelle coun s, a
de eca ion a e pe day was calcula ed o each coun
which was ob ained by di iding he numbe o pelle s
a each coun ing s a ion by he numbe o days since
he las coun a he same s a ion (Rouco e al., 2011).
Pelle pe sis ence can di e be ween habi a s and
seasons (Taylo & Williams, 1959; Ibo a & Luma e ,
1997; Paloma es, 2001; Fe nández de Simon e al.,
2011b) he e o e a 'pelle decay a e' was es ima ed o
ensu e an adequa e pe iod be ween pelle clea ance
and coun ing, ollowing he me hod o Fe nández de
Simon e al. (2011b).
Food a ailabili y
To quan i y ood abundance, a second pe manen plo
(1 × 1 m) was placed a each o he 146 sampling
poin s. To al g een ege a ion co e and co e pe
plan species we e es ima ed seasonally ollowing he
me hod p oposed by B aun–Blanque (1979). G een
co e was chosen o quan i y ood a ailabili y because
g azing species will ac i ely selec g een pa s o plan s
om a ela i ely d y swa d du ing d y seasons (Ja �
man & Sinclai , 1979). A high numbe o he baceous
species we e iden i ied du ing sampling, so o educe
he numbe o a iables, we g ouped species by am�
ily. Thus, a o al o 32 amilies we e conside ed in he
analysis (Gallego Fe nández e al., 2006).
Re uge a ailabili y
Wa en densi y was assessed only in sp ing, in a 3–ha
ci cula plo placed a ound each sampling poin
(N = 146). Due o he leng h o ou s udy, we assumed
ha abbi densi y did no change. A ci cula plo o
3 ha was selec ed because i is simila o he la ges
home ange obse ed in Eu opean abbi popula ions
in a nea by a ea (Fe nández, 2005). Each wa en
wi hin a ci cula plo was geo– e e enced and he
dis ance be ween sampling poin and he nea es
wa en was measu ed using A cGIS 9.3.
Sc ub co e (mainly R. monospe ma) was es i�
ma ed as a pe cen age o co e in he 3–ha ci cula
plo placed a ound each sampling poin . Pe cen age
co e was assessed by using A cGIS 9.3 o digi alize
each R. monospe ma bush om ae ial pho og aphs
(scale 1:5,000) aken in 2001.
Da a analysis
We used pai wise co ela ions o es o co–linea i y
be ween he explana o y a iables (i.e. pe cen age
o al g een co e and pe cen age he b co e by
each amily, wa en densi y, dis ance o nea es
wa en, and pe cen age R. monospe ma co e ). As
no case o co–linea i y was obse ed, all a iables
we e conside ed in he model. In o de o de ine
Fig. 1. Geog aphical loca ion o he s udy a ea and Re ama monospe ma dis ibu ion (in black).
Fig. 1. Localización geog á ica de la zona de es udio y dis ibución de Re ama monospe ma (en neg o).
La An illa
ou is u baniza ion
Ma shes
Pied as Ri e Es ua y
A lan ic Ocean 1 km
Spain
37º 12' N
7º 10' W
236 Della io e e al.
habi a p e e ence by abbi s a ou s udy si e, a
mul iple eg essions analysis pe season was used
o gene a e models ha included all ou explana o y
a iables as p edic o s, and abbi ela i e abundance
as ou dependen a iable. Using a model selec ion
app oach based on in o ma ion heo y (Bu nham
& Ande son, 2002) we iden i ied he se o models
bes suppo ed by he da a. The Akaike in o ma ion
c i e ion (AIC) was used o ank and weigh compe ing
models, using he di e ence be ween AICs o each
model and he minimum AIC ound (ΔAIC) (Bu nham
& Ande son, 2002). These ΔAIC alues we e also
used o calcula e Akaike’s weigh ing o each model wi
(Bu nham & Ande son, 2002). The alues o wi we e
s anda dized ac oss he candida e se o models.
Weigh ings also p o ide a way o de ine he ela i e
impo ance o each p edic o . This measu e can be
calcula ed as he sum o wi alues o e all he models
ha include he p edic o o in e es . Howe e , i is
i ually impossible o disce n he ela i e in luence
o di e en p edic o s when hey all appea in he
bes se o models, so we used he s anda dized
eg ession coe icien s o ank he impo ance o he
p edic o s in he bes model.
All s a is ical analyses we e pe o med using S a�
is ica (S a So , Inc., 2003) and Spa ial Analyses in
Mac oecology (SAM) (Rangel e al., 2005).
Resul s
Rabbi ela i e abundance
Mean aecal pelle pe sis ence was 0.99 and no di�
e ences we e obse ed be ween seasons. A e age
pelle de eca ion a e pe day pe si e was simila
h oughou he yea 0.71 (+ 0.015), 0.64 (+ 0.010),
0.98 (+0.021), 0.55 (+0.014) in win e , sp ing, summe
and au umn, espec i ely; howe e , signi ican di e�
ences we e obse ed be ween summe and au umn
(Z = –1.6; P = 0.052).
A new species o Laemos enus Bonelli, 1810
(Coleop e a, Ca abidae) om Els Po s Na u al Pa k
(Ca alonia, no heas e n Ibe ian peninsula)
Summe models
Fo summe we ob ained 11 models wi h ∆AIC < 2
and each explaining a simila amoun o a iance
(app ox. 20%) ( able 2). Wa en densi y a ia�
bles we e p esen in all models. Howe e , he
Akaike weigh s sugges ed ha he bes model
includes wa en densi y and R. monospe ma co e
(wi = 0.087). The highe s anda dized coe icien s o
bo h a iables p o ided s ong e idence ha hese
a e he main ac o s d i ing habi a use by abbi s
in summe a El Rompido spi .
Au umn models
Fo au umn, ou equi alen models we e ob ained
wi h ∆AIC < 2 and each explained a simila amoun
o a iance (app ox. 92%) ( able 3). R. monospe ma
co e and wa en densi y we e common p edic o s
o all models. Howe e , he Akaike weigh ings sug�
ges ed ha he bes model included wa en densi y,
R. monospe ma co e and co e o he bs o he
Ge aniaceae amily (nega i e) (wi = 0.272). The
highe s anda dized coe icien s o wa en densi y
and R. monospe ma co e p o ided s ong e idence
ha hese wo a iables we e he main ac o s d i ing
habi a use by abbi s in au umn.
Table 1. En i onmen al uni s on he 'La Flecha de El Rompido' s udy a ea, SW Spain.
Tabla 1. Unidades ambien ales en la zona de es udio de la Flecha del Rompido, SO España.
En i onmen al uni s A ea (ha) % Sample poin s
Dune sc ubland 26.6 5.06 10
R. monospe ma wi h Thymus ca nosus 72.8 13.84 18
R. monospe ma wi h A emisia c i hmi olia 112.3 21.44 26
R. monospe ma wi h Helich ysum pica dii 46.9 8.92 13
Inland dunes wi h R. monospe ma 11.4 2.17 10
Ac i e dune a ea 61.9 11.77 16
Tidal swales 93.4 17.76 20
Sal ma shes 32.7 6.22 10
Sandy sho e 46.0 8.74 13
We dep ession 21.4 4.07 10
S udy a ea o al 522.4 100 146
Animal Biodi e si y and Conse a ion 37.2 (2014) 237
Win e models
Fo win e , ou equi alen models we e ob ained wi h
∆AIC < 2, each explaining a simila amoun o a iance
(app ox. 37%) ( able 4). Fou a iables we e common
o all models: wa en densi y, R. monospe ma co e ,
co e o he bs o he U icaceae amily and o al he b
co e (nega i e). The Akaike weigh ings sugges ed
ha he bes model included he ou a iables men�
ioned abo e (wi = 0.314). The highe s anda dized
coe icien s o wa en densi y, R. monospe ma co e
and co e o U icaceae he bs p o ided s ong e i�
dence ha hese h ee a iables we e he main ac o s
d i ing habi a use by abbi s in win e .
Sp ing models
Fo sp ing, 10 models we e ob ained wi h ∆AIC < 2 each
explaining a simila amoun o a iance (app ox. 40%)
( able 5). Co e o he bs o he Bo aginaceae amily,
wa en densi y and R. monospe ma co e we e p esen
in all models. The Akaike weigh ings sugges ed ha
he bes model included he h ee a iables men ioned
abo e plus o al he b co e (nega i e), co e o Cheno�
podiaceae he bs (nega i e) and co e o Ge aniaceae
he bs (nega i e) (wi = 0.140). The highe s anda dized
coe icien s o co e o he bs o he Bo aginaceae am�
ily, wa en densi y, and R. monospe ma co e p o ide
s ong e idence ha hese h ee a iables a e he main
ac o s d i ing habi a use by abbi s in sp ing.
Discussion
Coas al dune sys ems wi h we dep essions ha e been
conside ed a a ou able habi a o wild abbi s because
we a eas would p o ide su icien ood esou ces
(Paloma es, 2003; Gál ez–B a o, 2011) and allow he
animals o build la ge wa ens (Paloma es, 2003). O he
au ho s ha e also ound his en i onmen al uni was
ha mos commonly used by abbi s o build wa ens
(Della io e e al., 2008). The explana ion o his ind�
ing could be he e y high co e o R. monospe ma
(90–100%) in his ecosys em (Vallés e al., 2011),
in e wined wi h all old sh ubs and p o iding excellen
p o ec ion ( o abbi s and wa ens) agains p eda o s
(Delibes–Ma eos e al., 2008a). R. monospe ma also
appea s o be an impo an sou ce o ood o abbi s
(Gómez Sal e al., 1999; Della io e e al., 2006). I is
no he e o e su p ising ha R. monospe ma has been
ound o be selec ed by wild abbi s independen ly o
season in mos well–suppo ed models.
Table 2. Mul iple eg ession models o Eu opean abbi s in summe . Models a e anked by AIC om
bes – o wo s – i ing, and only models wi h ∆AIC < 2 a e p esen ed. AICs ha e been co ec ed o
he p esence o spa ial au oco ela ion in he model esiduals. Fo each a iable en e ing he model
we included i s s anda dized coe icien o e alua e i s ela i e impo ance: Wd. Wa en densi y; Rm.
Re ama monospe ma; Chen. Chenopodiaceae; Plum. Plumbaginaceae.
Tabla 2. Modelos de eg esión múl iple pa a el conejo de mon e en e ano. Los modelos es án o denados
del mejo al peo alo de AIC. Solo p esen amos modelos cuyo ∆AIC < 2. Los alo es de AIC de
los modelos es an es se co igie on pa a ene en cuen a la exis encia de au oco elación espacial.
Pa a cada a iable del modelo incluimos un coe icien e es anda izado pa a e alua su impo ancia
ela i a: Wd. Densidad de mad igue as; Rm. Re ama monospe ma; Chen. Chenopodiaceae; Plum.
Plumbaginaceae.
Rm Chen Co e Plum
Wd co e (%) co e (%) o he bs (%) co e AIC ∆AIC 2 wi
0.394 0.151 1237.6 0 0.208 0.087
0.395 0.154 −0.105 1237.7 0.1 0.220 0.082
0.386 0.160 −0.092 1238.1 0.5 0.218 0.068
0.140 1238.1 0.5 0.193 0.066
0.441 −0.102 1238.5 0.9 0.203 0.055
0.405 0.134 −0.09 1238.6 1 0.215 0.053
0.438 −0.10 1238.6 1 0.203 0.053
0.388 0.163 −0.09 −0.074 1238.8 1.2 0.226 0.047
0.436 −0.084 1238.1 1.5 0.200 0.042
0.398 0.148 −0.08 −0.077 1239.4 1.8 0.222 0.035
0.402 0.145 −0.083 −0.059 1239.5 1.9 0.222 0.033

238 Della io e e al.
species o his amily a e annuals, ha lowe and
ui om Feb ua y o June, and a e p esen only in
en i onmen al uni s wi h R. monospe ma and dune
sc ubland. Bo h amilies could be associa ed wi h
abbi habi a selec ion because bo h a e pala able
o abbi s (Della io e, 2007; Della io e e al., 2009)
and ha e a highe occu ence on inland dunes wi h
R. monospe ma, which is he habi a selec ed by
abbi s o wa en building (Della io e e al., 2008).
On he o he hand, o al he b co e and Chenopo�
diaceae and/o Ge aniaceae co e we e nega i ely
associa ed wi h abbi p esence du ing au umn, win e
Table 3. Mul iple eg ession models o Eu opean abbi s in au umn. Models a e anked by AIC om bes –
o wo s – i ing, and only models wi h ∆AIC < 2 a e p esen ed. AICs ha e been co ec ed o he p esence
o spa ial au oco ela ion in he model esiduals. Fo each a iable en e ing he model, we included i s
s anda dized coe icien o e alua e i s ela i e impo ance: Rm. Re ama monospe ma; Wd. Wa en densi y;
Ge . Ge aniaceae; Ama . Ama yllidaceae.
Tabla 3. Modelos de eg esión múl iple pa a el conejo de mon e en o oño. Los modelos es án o denados del
mejo al peo alo de AIC. Solo p esen amos modelos cuyo ∆AIC < 2. Los alo es de AIC de los modelos
es an es se co igie on pa a ene en cuen a la exis encia de au oco elación espacial. Pa a cada a iable del
modelo incluimos un coe icien e es anda izado pa a e alua su impo ancia ela i a: Rm. Re ama monospe ma;
Wd. Densidad de mad igue as; Ge . Ge aniaceae; Ama . Ama yllidaceae.
Rm Ge Ama
co e (%) Wd co e (%) co e (%) AIC ∆AIC 2 wi
0.678 0.476 −0.119 721.9 0 0.926 0.272
0.628 0.482 722.2 0.3 0.924 0.237
0.704 0.445 −0.096 722.9 0.9 0.925 0.172
0.757 0.452 −0.107 −0.073 723.4 1.5 0.926 0.132
Table 4. Mul iple eg ession models o Eu opean abbi s in win e . Models a e anked by AIC om bes – o
wo s – i ing, and only models wi h ∆AIC < 2 a e p esen ed. AICs ha e been co ec ed o he p esence
o spa ial au oco ela ion in he model esiduals. Fo each a iable en e ing he model we included i s
s anda dized coe icien o e alua e i s ela i e impo ance: Wd. Wa en densi y; Rm. Re ama monospe ma;
U ic. U icaceae; P im. P imulaceae; Plum. Plumbaginaceae.
Table 4. Modelos de eg esión múl iple pa a el conejo de mon e en in ie no. Los modelos es án o denados
del mejo al peo alo de AIC. Solo p esen amos los modelos cuyo ∆AIC < 2. Los alo es de AIC de los
modelos es an es se co igie on pa a ene en cuen a la exis encia de au oco elación espacial. Pa a cada
a iable del modelo incluimos un coe icien e es anda izado pa a e alua su impo ancia ela i a: Wd. Densidad
de mad igue as; Rm. Re ama monospe ma; U ic. U icaceae; P im. P imulaceae; Plum. Plumbaginaceae.
Rm U ic He b. P im Plum
Wd co e (%) co e (%) co e (%) co e (%) co e (%) AIC ∆AIC 2 wi
0.293 0.258 0.213 −0.172 1185.6 0 0.373 0.314
0.291 0.259 0.217 −0.174 0.079 1186.4 0.8 0.379 0.213
0.299 0.255 0.206 −0.172 –0.073 1186.6 1 0.378 0.187
0.303 0.254 0.209 −0.176 0.079 –0.069 1187.6 2 0.384 0.118
Al hough we obse ed some seasonal di e ences
be ween he a iables ha p edic ed habi a use by
abbi s ( ables 2–5), ou esul s showed ha wa en
densi y and R. monospe ma co e we e he main
p edic o s h oughou he whole yea . The di e ences
obse ed be ween seasons could be due o changes
in ege a ion a ailabili y. Fo example, Bo aginaceae
co e appea ed as a likely p edic o o abbi habi a
use in sp ing because he species o his amily a e
annuals ha lowe and ui in sp ing; and g ow ex�
clusi ely on inland dunes wi h R. monospe ma. The
same applied o he U icaceae amily in win e . The
Animal Biodi e si y and Conse a ion 37.2 (2014) 239
Table 5. Mul iple eg ession models o Eu opean abbi s in sp ing. Models a e anked by AIC om
bes – o wo s – i ing, and only models wi h ∆AIC < 2 a e p esen ed. AICs ha e been co ec ed
o he p esence o spa ial au oco ela ion in he model esiduals. Fo each a iable en e ing he
model we included i s s anda dized coe icien o e alua e i s ela i e impo ance: Bo . Bo aginaceae;
Wd: Wa en densi y; Rm. Re ama monospe ma; Chen. Chenopodiaceae; Ge . Ge aniaceae; P im.
P imulaceae.
Tabla 5. Modelos de eg esión múl iple pa a el conejo de mon e en p ima e a. Los modelos es án o denados
del mejo al peo alo de AIC. Solo p esen amos los modelos cuyo ∆AIC < 2. Los alo es de AIC de los
modelos es an es se co igie on pa a ene en cuen a la exis encia de au oco elación espacial. Pa a
cada a iable del modelo incluimos un coe icien e es anda izado pa a e alua su impo ancia ela i a:
Bo . Bo aginaceae; Wd. Densidad de mad igue as; Rm. Re ama monospe ma; Chen. Chenopodiaceae;
Ge . Ge aniaceae; P im. P imulaceae.
Rm Chen He b Ge
Bo Wd co e (%) co e (%) co e (%) co e (%) P im AIC ∆AIC 2 wi
0.346 0.304 0.276 −0.129 −0.115 −0.115 1030.2 0 0.419 0.140
0.339 0.314 0.267 −0.136 −0.119 1030.4 0. 8 0.409 0.127
0.343 0.316 0.285 −0.132 −0.117 −0.120 0.071 1031 0.8 0.425 0.094
0.348 0.292 0.267 −0.12 −0.114 1031.4 1.2 0.406 0.092
0.343 0.305 0.105 −0.125 1031.6 1.4 0.394 0.070
0.338 0.327 0.272 −0.139 −0.119 0.06 1031.8 1.6 0.412 0.062
0.356 0.290 0.274 −0.111 −0.129 0.083 1032 1.8 0.412 0.062
0.352 0.289 0.264 −0.119 1032. 1.8 0.393 0.056
0.347 0.302 0.277 −0.114 −0.100 1032. 1.8 0.401 0.054
and sp ing ( ables 2–5). Species o Chenopodiaceae
a e abundan in idal swales, sal ma shes and on sandy
sho es whe eas species o Ge aniaceae a e abundan
on ac i e dunes and in dune sc ubland. The lack o
shel e o p eda o s in hese en i onmen al uni s could
explain why abbi s did no selec hese uni s.
O he au ho s ound seasonal di e ences in habi a
use by abbi s in g assland en i onmen s (Rueda
Ga cía, 2006; Rueda e al., 2008a, 2008b). In ou
s udy a ea, al hough we did no es o di e ences
in habi a use be ween seasons, we ound no change
in habi a use in each season. This may indica e ha
ood a ailabili y in he icini y o wa ens emains
ela i ely cons an o e ime; hence abbi s do no
need o mo e long dis ances o eeding. G assland
and he baceous ege a ion biomass in ou s udy a ea
dec ease du ing summe due o he high empe a u es
and low ain all. Howe e , eeding si es selec ed by
abbi s emained cons an and we e always co ela ed
wi h wa en densi y and R. monospe ma co e . Thus,
abbi s may eed almos en i ely on such ui s un il
la e au umn; in win e , when hose ui s become
sca ce, abbi s eed on he baceous ege a ion, which
is highly a ailable due o ain all in ha season. Ac�
co dingly, when he baceous ege a ion is sca ce,
ui a ailabili y is high and abbi s can eed in a eas
p o ec ed om p eda o s.
Al hough he densi y o indi iduals migh be a poo
indica o o habi a quali y in some condi ions (Van
Ho ne, 1983; Ly a–Jo ge e al., 2010), mos ly i is a
good p oxy o he sui abili y (i.e. quali y) o a pa �
icula a ea. Fo Eu opean abbi popula ions, he key
limi ing ac o s a e he inabili y o dig wa ens and he
lack o a ailable e uges and ood (Roge s & Mye s,
1979; Paloma es & Delibes, 1997; Fa e al., 1999;
Lomba di e al., 2003; Vi gos e al., 2003; Cab e a
Rod íguez, 2006). In coas al dune sys ems, whe e
he e ain is i m sand, wa en cons uc ion is no a
limi ing ac o (Moseby e al., 2005). Food a ailabili y
should no be a limi ing ac o ei he conside ing he
high p esence o R. monospe ma ui s, which a e
highly consumed by abbi s in ou s udy si e (Del�
la io e e al., 2006). Re uge a ailabili y, o p o ec ion
agains p eda o s, howe e , may be he key ac o o
his pa icula popula ion. A p e ious s udy showed
ha Eu opean abbi s a ou s udy si e equi ed high
co e o R. monospe ma (90–100%) o wa en build�
ing (Della io e e al., 2008). This would occu mo e
likely in sand dune a eas whe e he e is no s uc u al
suppo (i.e. no sh ub oo s), making hem unsui able
o wa en building (e.g. Ma ins e al., 2002; Palo�
ma es, 2003). R. monospe ma also p o ides dense
o e head co e and acili a es he es ablishmen and
g ow h o o he plan species, mainly he baceous
annuals, unde he sh ub canopies. This ege a ion
would also p o ide ood o abbi s (Muñoz–Vallés
e al., 2014), cha ac e is ics ha a e co ela ed wi h
abbi occu ence and ela i e abundance in sc ub�
240 Della io e e al.
land habi a s (Ma ins e al., 2002; Beja e al., 2007).
Un o una ely, p eda o p esence was no moni o ed
a ou s udy si e, bu p eda o s such as he ed ox
(Vulpes ulpes) a e qui e common in he a ea and
hey may a ec he iabili y o he abbi popula ions,
since hey could ac as acul a i e p eda o o abbi s
(Delibes–Ma eos e al., 2008b).
In conclusion, ou esul s show ha in a coas al
sand dunes ecosys em in he sou h–wes e n Ibe ian
Peninsula, sc ub co e , and in pa icula R. monos-
pe ma co e , is highly p e e ed by Eu opean abbi s,
independen ly o season. Al hough R. monospe ma
has ecen ly been ca ego ized as a na i e–in asi e
plan in coas al dune sys ems (Muñoz–Vallés e al.,
2014), he species seems o bene i wild abbi popula�
ions in such ecosys ems. I p o ides a comp ehensi e
habi a o abbi s: on he one hand, i s canopy p o i�
des p o ec ion agains p eda o s (e.g. agains ae ial
p eda o s) and i s oo sys em p o ides s uc u e o
he soil o enable wa en building, while on he o he
hand, i p o ides ood by mean o i s ui s and by
a ou ing he baceous g ow h unde nea h. The e o e,
he bene i s o R. monospe ma should be aken in o
accoun when deciding on u u e managemen s a�
egies o wild abbi s. Fo example, abbi s in coas al
dunes could be used as a po en ial popula ion sou ce
o inc ease abbi numbe s o conse a ion pu poses
in adjacen habi a s.
Acknowledgemen s
This pape epo s Ph. D. esea ch by CD a he
Uni e si y o Se ille (Spain). CD and CR con ibu ed
equally o his pa icula wo k. We hank he s a o he
Pa aje Na u al Ma ismas del Río Pied as y Flecha de
El Rompido o allowing us o conduc esea ch a he
si e. Family, s uden s, and iends gene ously helped
wi h ieldwo k. We also hank D Isabel C Ba io, D Luis
Tapia and he ABC Edi o o hei help ul commen s on
p e ious d a s o he manusc ip , and Ch is ine Beza
om Landca e Resea ch o he edi o ial commen s
in he manusc ip .
Re e ences
Ba io, I. C., Ace edo, P. & To osa, F. S., 2009. As�As�
sessmen o me hods o es ima ing wild abbi
popula ion abundance in ag icul u al landscapes.
Eu opean Jou nal o Wildli e Resea ch, 56: 335–340.
Beja, P., Pais, M. & Palma, L., 2007. Rabbi O yc ola-
gus cuniculus habi a s in Medi e anean sc ubland:
he ole o sc ub s uc u e and composi ion. Wildli e
Biology, 13: 28–37.
Bond, B. T., Bu ge , L. W., Leopold, B. D., Jones, J.
C. & Godwin, K. D., 2002. Habi a use by co on ail
abbi s ac oss mul iple spa ial scales in Mississippi.
Jou nal o Wildli e Managemen , 66: 1171–1178.
Bos, D. G. & Ca hew, S. M., 2003. The in luence o
beha iou and season on habi a selec ion by a
small mammal. Ecog aphy, 26: 810–820.
B aun–Blanque , J., 1979. Fi osociología: bases
pa a el es udio de las comunidades ege ales. H.
Blume, Mad id.
Bu nham, K. P. & Ande son, D. R., 2002. Model selec ion
and mul imodel in e ence: a p ac ical in o ma ion–
heo e ic app oach, 2nd edi ion. Sp inge , New Yo k.
Cab e a Rod íguez, F., 2006. Mic ohabi a selec ion o
he Eu opean abbi on La Palma, Cana y Island,
Spain. Ac a The iologica, 51: 435–442.
Delibes–Ma eos, M., Delibes, M., Fe e as, P. & Vil�
la ue e, R., 2008a. Key ole o Eu opean abbi s in
he conse a ion o he wes e n Medi e anean ba�
sin ho spo . Conse a ion Biology, 22: 1106–1117.
Delibes–Ma eos, M., Fe nández de Simon, J., Villa�
ue e, R. & Fe e as, P., 2008b. Feeding esponses
o he ed ox (Vulpes ulpes) o di e en wild
abbi (O yc olagus cuniculus) densi ies: a egional
app oach. Eu opean Jou nal o Wildli e Resea ch,
54: 71–78.
Della io e, C. M., 2007. Ecología del conejo sil es�
e (O yc olagus cuniculus) en un sis ema duna .
Disse a ion, Uni e sidad de Se illa.
Della io e, C. M., Gallego Fe nández, J. B. & Vallés,
S. M., 2008. Habi a use o wa en building by
Eu opean abbi s (O yc olagus cuniculus) in ela�
ion o landscape s uc u e in a sand dune sys em.
Ac a Oecologia, 33: 372–379.
– 2009. The abbi (O yc olagus cuniculus) as a seed
dispe se in a coas al dune sys em. Plan Ecology,
206: 251–261.
Della io e, C. M., Vallés, S. M. & Gallego Fe nández,
J. B., 2006. Rabbi s (O yc olagus cuniculus) as
dispe se s o Re ama monospe ma (L.) seeds
in a coas al dune sys em. Ecoscience, 13: 5–10.
Fa, J. E., Sha ples, C. M. & Bell, D. J., 1999. Habi a
co ela es o Eu opean abbi (O yc olagus cuniculus)
dis ibu ion a e he sp ead o RVHD in Cadiz
P o ince, Spain. Jou nal o Zoology, 249: 83–96.
Fe nández, N., 2005. Spa ial pa e ns in Eu opean
abbi abundance a e a popula ion collapse.
Landscape Ecology, 20: 897–910.
Fe nández de Simon, J., Díaz–Ruiz, F., Ci illi, F., To �
osa, F. S., Villa ue e, R., Delibes–Ma eos, M. &
Fe e as, P., 2011a. Towa ds a s anda dized index
o Eu opean abbi abundance in Ibe ian Medi�
e anean habi a s. Eu opean Jou nal o Wildli e
Resea ch, 57: 1091–1100.
Fe nández de Simon, J., Díaz–Ruiz, F., Villa ue e,
R., Delibes–Ma eos, M. & Fe e as, P., 2011b. As�As�
sessing p edic o s o pelle pe sis ence in Eu opean
abbi s O yc olagus cuniculus: owa ds eliable
popula ion es ima es om pelle coun s. Wildli e
Biology, 17: 317–325.
Gál ez–B a o, L. 2011. Conejo –O yc olagus cuni-
culus. En: Enciclopedia Vi ual de los Ve eb ados
Españoles (A. Sal ado & J. Cassinello, Eds.).
Museo Nacional de Ciencias Na u ales, Mad id.
h p://www. e eb adosibe icos.o g/
Gallego Fe nández, J. B., Vallés, S. M. & Della io e,
C. M., 2006. Nue a Umb ia Spi . Flo a and ege-
a ion. Ed. Exmo Ayun amien o de Lepe. Huel a.
Gómez Sal, A., Rey Benayas, J. M., Pin o , L. & Re�
bollo, S., 1999. Role o dis u bance in main aining
a sa anna–like pa e n in Medi e anean Re ama
Animal Biodi e si y and Conse a ion 37.2 (2014) 241
sphae oca pa sh ubland. Jou nal o Vege a ion
Science, 10: 365–370.
Ha gis, C. D. & McCullough, D. R., 1984. Win e
die and habi a selec ion o Ma en in Yosemi e
Na ional Pa k. Jou nal o Wildli e Managemen ,
48: 140–146.
He nandez–Aguila , R. A., Moo e, J. & S an o d, C.
B., 2013. Chimpanzee nes ing pa e ns in sa anna
habi a : En i onmen al in luences and p e e ences.
Ame ican Jou nal o P ima ology, 75: 979–994.
Ibo a, O. & Luma e , J. P., 1997. Validi y limi s o
he pelle g oup coun s in wild abbi (O yc olagus
cuniculus). Mammalia, 61: 205–218.
Jaksic, F. M., Seib, R. L. & He e a, C. M., 1982.
P eda ion by he ba n owl (Ty o alba) in Medi e �
anean habi a s o Chile, Spain and Cali o nia: A
compa a i e app oach. Ame ican Midland Na u al-
is , 107: 151–162.
Jaksic, F. M. & So igue , R. C., 1981. P eda ion upon
he Eu opean Rabbi (O yc olagus cuniculus) in
Medi e anean habi a s o Chile and Spain – A
compa a i e–analysis. Jou nal o Animal Ecology,
50: 269–281.
Ja man, P. J. & Sinclai , A. R. E., 1979. Feeding s a egy
and he pa e n o esou ce pa i ioning in A ican
ungula es. In: Se enge i: dynamics o an ecosys em:
130–164 (A. R. E. Sinclai & M. No on–G i i hs,
Eds.). Uni e si y o Chicago P ess, Chicago.
Ley, C., Gallego–Fe nández, J. B. & Vidal, C., 2011.
Coas al dunes es o a ion manual. Minis e io
de Medio Ambien e, Ru al y Ma ino, San ande ,
España.
Lisón, F., Haz, Á. & Cal o, J. F., 2014. Habi a p e �Habi a p e �
e ence o he me idional se o ine ba Ep esicus
isabellinus (Temminck, 1840) in semia id Medi e �
anean landscapes. Animal Biodi e si y and Con-
se a ion, 37.1: 59–67.
Lomba di, L., Fe nández, N. & Mo eno, S., 2007.
Habi a use and spa ial beha iou in he Eu opean
abbi in h ee Medi e anean en i onmen s. Basic
and Applied Ecology, 8: 453–463.
Lomba di, L., Fe nández, N., Mo eno, S. & Villa ue e,
R., 2003. Habi a – ela ed di e ences in abbi
(O yc olagus cuniculus) abundance, dis ibu ion
and ac i i y. Jou nal o Mammalogy, 84: 26–36.
Ly a–Jo ge, M. C., Ribei o, M. C., Cioche i, G.,
Tambosi, L. R. & Pi ello, V. R., 2010. In luence o
mul i–scale landscape s uc u e on he occu ence
o ca ni o ous mammals in a human–modi ied
sa anna, B azil. Eu opean Jou nal o Wildli e
Resea ch, 56: 359–368.
Ma ins, H., Mille , D. R., Els on, D. A., Rego, F. &
Milne, J. A., 2002. Fac o s in luencing he loca ion
and numbe o en ances o Eu opean wild abbi
(O yc olagus cuniculus L.) wa ens in a Sou he n
Po uguese mon ado. Jou nal o Medi e anean
Ecology, 3: 31–40.
Mo ison, M. L., Ma co , B. G. & Mannan, R. W., 2006.
Wildli e habi a ela ionships: concep s and applica-
ions. Thi d edi ion. Island P ess, Washing on, DC.
Moseby, K. E., De Jong, S., Mun o, N. & Pieck,
A., 2005. Home ange, ac i i y and habi a use
o Eu opean abbi s (O yc olagus cuniculus) in
a id Aus alia: implica ions o con ol. Wildli e
Resea ch, 32: 305–311.
Muñoz–Vallés, S., Gallego–Fe nández, J. B. & Cam�
b ollé, J., 2014. The ole o he expansion o na�
i e–in asi e plan species in coas al dunes: The
case o Re ama monospe ma in SW Spain. Ac a
Oecologia, 54: 82–89.
Mye s, K., 1955. Cop ophagy in he Eu opean ab�
bi s (O ycolagus cuniculus) in Aus alia. Aus alian
Jou nal o Zoology, 3: 336–345.
Paloma es, F., 2001. Compa ison o 3 me hods o
es ima e abbi abundance in a Medi e anean
en i onmen . Wildli e Socie y Bulle in, 29: 578–585.
– 2003. Wa en building by Eu opean abbi s
(O yc olagus cuniculus) in ela ion o co e a ail�
abili y in a sandy a ea. Jou nal o Zoology, London,
259: 63–67.
Paloma es, F. & Delibes, M., 1997. P eda ion upon Eu o�P eda ion upon Eu o�
pean abbi s and hei use o open and closed pa ches
in Medi e anean habi a s. Oikos, 80: 407–410.
Pulliainen, E. & Tunkka i, P., 1987. Win e die , habi a
selec ion and luc ua ion o a moun ain ha e Le-
pus imidus popula ion in Finnish Fo es Lapland.
Hola c ic Ecology, 10: 261–267.
Rangel, T. F. L. V. B., Diniz–Filho, J. A. F. & Bini, L.
M., 2005. SAM .1. 0 be a – Spa ial Analysis in
Mac oecology (So wa e and Use ’sGuide) (h p://
www.Ecoe ol.u g.b /sam/index.h m).
Roge s, P. M. & Mye s, K., 1979. Ecology o he
Eu opean wild abbi , O yc olagus cuniculus L.
in Medi e anean habi a s. I. Dis ibu ion in he
landscape o he Co o Doñana, S. Spain. Jou nal
o Applied Ecology, 16: 691–703.
Rouco, C., Villa ue e, R., Cas o, F. & Fe e as, P.,
2011. Responses o naï e and expe ienced Eu o�
pean abbi s o p eda o odou . Eu opean Jou nal
o Wildli e Resea ch, 57: 395–398.
Rueda Ga cía, M., 2006. Selección de hábi a po
he bí o os de di e en e amaño y sus e ec os
sob e la ege ación. El papel del conejo eu opeo
(O yc olagus cuniculus) en ecosis emas de dehesa.
Disse a ion, Uni e sidad de Alcalá.
Rueda, M., Rebollo, S. & Gál ez B a o, L., 2008a.
Age and season de e mine Eu opean abbi habi a
use in Medi e anean ecosys em. Ac a Oecologia,
34: 266–273.
Rueda, M., Rebollo, S., Gál ez B a o, L. & Escude o,
A., 2008b. Habi a use by la ge and small he bi�Habi a use by la ge and small he bi�
o es in a luc ua ing Medi e anean ecosys em:
Implica ion o seasonal changes. Jou nal o A id
En i onmen s, 72: 1698–1708.
Sa men o, P., C uz, J., Paula, A., Ei a, C., Capinha,
M., Amb ósio, I., Fe ei a, C. & Fonseca, C., 2012.
Occupancy, coloniza ion and ex inc ion pa e ns
o abbi popula ions: Implica ions o Ibe ian lynx
conse a ion. Eu opean Jou nal o Wildli e Re-
sea ch, 58: 523–533.
San illi, F. & Bagliacca, M., 2010. Habi a use by he
Eu opean wild abbi (O yc olagus cuniculus) in
a coas al sandy dune ecosys em o Cen al I aly.
Hys ix – I alian Jou nal o Mammology, 21: 57–64.
S o , P., 2003. Use o space by sympa ic Eu opean
ha es (Lepus eu opaeus) and Eu opean abbi s