Hea y looding e ec s on home ange and
habi a selec ion
o
ee- anging Ibe ian
ha es
(
Lepus g ana ensis
)
in
Doña
na
Na ional Pa k (SW
Spain)
F ancisco Ca o
&
Ramón C. So igue
&
J. F. Bel án
&
Ana C.
And eu
Abs ac The Ibe ian ha e Lepus g ana ensis is a common
and abundan species h oughou he Ibe ian Peninsula;
howe e , s udies documen ing i s ecology a e sca ce.
Be ween Augus 1996 and Sep embe 1997, a e a long
d ough , we s udied he e ec s o a widesp ead lood on he
Ibe ian ha e ecology in eco one pas u es o Doñana
Na ional Pa k. Ha es we e cap u ed using e ical ne s
h ough pa o 1996 and 1997. Fou een adul ha es (nine
emales and i e males) we e agged wi h adiocolla s and
loca ed by iangula ion on a daily basis a leas once a
week. Habi a use was es ima ed using a geog aphical
in o ma ion sys em. The a e age home ange size was
28 ha o males and 24 ha o emales. No signi ican
di e ences we e obse ed be ween sexes. Signi ican
di e ences we e obse ed in he use o sc ub a eas and in
d y pas u es. Du ing he d y and we seasons, males and
emales we e mos o en loca ed in he pas u es o he
eco one,
ush s ands and sc ub. The dense b acken (P e idium
aquilinum) and seaside bul ush (Sci pus ma i imus) commu-
ni ies we e a oided. These esul s a e he i s on he spa ial
ecology o his species in i s na i e habi a .
C
ommunica ed by: Ma W. Haywa d
F. Ca o
(
*
)
:
R. C.
So igue
:
A. C. And eu
Es ación Biológica de Doñana (C. S. I. C.),
A da Amé ico Vespucio s/n, Isla de la Ca uja,
41092 Se illa, Spain
e-mail: pca [email protected]
J. F. Bel án
Depa men o Physiology and Zoology, Uni e si y o Se ille,
A da. Reina Me cedes, 6,
41012 Se illa, Spain
Keywo ds Lepus g ana ensis . Ibe ian ha e . Home
ange
.
Habi a
selec ion
. Ibe ian
Peninsula
.
Doñana
. Flooding
In oduc ion
The p esence o an animal and i s spa ial dis ibu ion is
de e mined by (a) esou ces dis ibu ion (Co ich 1976;
Emlen and O ing 1977), (b) he p esence o absence o
o he indi iduals o he same species (B own and O ians
1970), (c) he a ailabili y o app op ia e habi a (Johnson
1980; Thomas and Taylo 1990) and (d) he p esence and
loca ion o p eda o s (Es es 2005; Ripple and Besch a
2004). As he densi y o animals inc eases, dispu es occu
o e esou ces, which o ces animals o mo e owa ds
habi a s wi h ewe esou ces bu less compe i ion (F e well
and Lucas 1969). Howe e , qui e o en, spa ial displace-
men is p oduced when he p e e ences habi a disappea
(o a e s ongly educed). These e ec s a e obse ed a e a
g ea snow co e o a e a lood (B oekhuizen and
Maaskamp 1982; Hewson and Hinge 1990; Wol 1980).
In his pape , we quan i y he e ec s on he changes o
habi a use p oduced by haza dous clima ic e en s changes
like a lood in he Doñana Na ional Pa k (DNP). Knowledge
o home anges and habi a use in he Ibe ian Ha e is e y
limi ed (de la Calzada and Ma ínez 1994; Rod íguez e al.
1997), and no wo k has e e s udied habi a selec ion and
home ange in his endemic Ibe ian species. Ou aim was o
documen Ibe ian
ha
e
’
s
spa ial ecology, including habi a
use and habi a selec ion, in a na u al,
undis u bed,
no-hun ed
a ea o Doñana Na ional Pa k wi h special a en ion o he
looding e ec s.
Ma e ial and
me hods
S udy a ea
This s udy was pe o med in he Doñana Na ional Pa k
(SW Ibe ian
Peninsula;
Fig. 1). The clima e is Medi e anean
sub-humid wi h mild and
unp edic able
ainy win e s and ho
d y summe s. Gene ally, ain all is concen a ed o mid-
sp ing. A long d ough , wi h a mean annual ain all less han
402 mm/yea , was obse ed du ing he pe iod 1990–1995.
An in ense ainy season p oducing looding was obse ed
om 13 Decembe 1996 o 31 Ap il 1997 (Me eo ological
S a ion: Palacio de Doñana). The yea s 1996 o 1997 we e
conside ed e y we , wi h annual ain all (abo e 1,000 mm)
as ly exceeding he long- e m mean annual ain all o he
a ea (506 mm, pe iod 1970–1996; U diales 1997). In ac ,
du ing au umn 1996, ain all was he 30-yea eco d and was
ma ched by he nex
yea
’
s
ain all again.
The main s udy a ea occu ed in he eco one zone
be ween ma shland and sh ubland (Fig. 1). Two o he mos
ou s anding cha ac e is ics o his eco one a e he g ea
di e si y o animal species ha li e he e and he ac ha
in some ainy yea s, i becomes looded e y quickly. The
ege a ion o his zone is domina ed by he plan associa-
ions
Sci pe um
ma i imi and Galio palus is wi h Junce um
ma i imum.
Cap u e me hods and adioma king
Ha es we e cap u ed and adio- acked be ween Augus
1996 and Sep embe 1997. Ha es we e caugh a nigh by
using a ehicle o d i e hem in o long ne s. The sex o all
animals was iden i ied by hei ex e nal geni alia, and hey
we e weighed o de e mine hei ages (Ca o 2005). This
me hod o cap u e does no ha m he ha es, and many
animals can be cap u ed in a sho space o ime (Kei h e
al. 1968). A o al numbe o 14 ee- anging ha es ( i e
males and nine emales) we e adio- acked (Table 1).
Each ha e was ea -ma ked wi h indi idually iden i iable
numbe ed plas ic ags and i ed wi h a adio- acking colla
o ca. 30 g, less o 1.5% o body weigh o adul ha e,
equipped wi h an ac i i y senso (Wagene adiocolla s,
Ge many) wi h a li e expen ancy o 6 o 7 mon hs. Ha es
we e loca ed one o wo imes a day on day ime and
sun ise, a leas once a week, depending on he season.
Loca ions we e pe o med by iangula ing pai s o ixed
s a ions. The loca ion e o es ima ed om expe imen al
ials was >10 m. Fo he home- ange analyses, we selec ed
eleme y ixes sepa a ed by a leas 12 h, in o de o a oid
any au oco ela ion be ween consecu i e loca ions (Swiha
and Slade 1985a, b). An inc emen al a ea analysis was
ca ied ou plo ing he size o home anges s. numbe o
loca ions (Kenwa d and Hodde 1992). We ound ha 25
Fig. 1 S udy a ea in he Doñana Na ional Pa k
Colla
Sex
Age
Da e o cap u e
Da e o las posi ion
No. o loca ions
144.613
F
SA
12/08/1996
12/12/1997
31
144.764
F
A
08/08/1996
12/12/1997
42
144.564
F
SA
08/08/1996
14/12/1997
43
144.714
M
J
08/08/1996
09/06/1997
68
144.690
F
A
12/08/1996
31/03/1997
35
144.914
F
A
12/08/1996
01/04/1997
41
144.589
F
A
13/08/1996
20/01/1997
28
144.539a
F
A
12/08/1996
20/01/1997
47
144.739
F
SA
13/08/1996
23/09/1997
108
144.888
M
A
04/03/1997
09/06/1997
31
144.839
M
A
11/03/1997
28/07/1997
43
144.789
F
A
12/03/1997
20/08/1997
58
144.764b
M
A
12/03/1997
06/09/1997
54
Table 1 Examples o Ibe ian
Ha e adio agged in eco one o
Doñana Na ional Pa k
Colla emission equency; Da e
o cap u e da e o i ing wi h
adio colla ; Da e o las posi ion
150.080 M A 13/03/1997 28/09/1997 60
da
e
o
las
posi
i
on
a a
il
able
loca ions we e enough o ob aining a s able es ima e o he
size o an Ibe ian
ha e
’
s
home ange.
Home ange, habi a use and habi a selec ion
Da a we e p e-p ocessed wi h he p og amme Loca e II
e sion 1.3 (Pace 1990). Calcula ions o es ima ing he
minimum angula e o be ween he obse ed angles and
he es ima ion o he loca ion o he animal we e ca ied ou
by means o a maximum likelihood es ima o (Len h 1981).
The sizes o he
ha es
’
home anges we e es ima ed using
he p og amme RANGES V (Kenwa d and Hodde 1996),
he home anges as exp essed by he minimum con ex
polygon ha connec ed he mos a - lung poin s o he
a i hme ical a e age o localisa ions o each animal (Moh
and S ump 1966; Jen ich and Tu ne 1969), using 90% o
adio localisa ions he eby excluding pe iphe al ixes which
could be classed as occasional explo a o y excu sions, o
calcula e he size o he home ange o Ibe ian ha es (Dixon
and Chapman 1980; Tappe and Ba nes 1986). We used his
es ima o o acili a ing compa ison wi h o he da a on
ha es al eady published.
Habi a
use was
calcula ed
wi h a
geog aphical
in o ma ion
sys em (Whi e
and
Ga od
1990) using he p og ammes Id isi
(Cla k Labs., USA) and A c iew 3.1.
Andalusian
Go e nmen
1:16,000
scale ae ial maps om
1997 and 1998 we e also used.
Fi s , a digi al ae ial pho og aph was
geo- e e enced
using he
p og amme Id isi (Cla k
Labs. USA). Six
classes
o ege a ion
we e
iden i ied
(sc ub, ushes,
b acken,
eeds, d y pas u e and
pas u es in
“
La
Ve a”, eco one) and hen digi alised and
con e ed in o polygons. The pe cen age o
use was calcula ed
as he pe cen age o he home ange lying in each o he
di e en
habi a s; he
a ailabili y
pe cen age was aken as he
ela i e p esence o each habi a wi hin he o al su ace a ea.
In o de o es o signi ican di e ences be ween male
and emale home anges, he Mann–Whi ney U es was
applied. Habi a selec ion was es ablished by means o
I le
’
s
Index (I le 1961; Ga neau e al. 2008; Sido o ich
e al 2008), in which he choice o habi a by he species
was calcula ed by he o mula:
IES : % Use − % A ailabili y = % Use
þ
% A ailabili y:
The alue o
I le
’
s
Index a ies be ween +1, which
would ep esen he exclusi e use o ha habi a by ha es,
and −1, which would indica e a o al ejec ion o ha
habi a . A alue o 0 indica es a neu al selec ion o he
habi a in ela ion o i s pe cen age p esence in he whole
a ea.
Resu
l s
Home ange size
The home ange es ima ed wi h a minimum con ex polygon
using 90% o adio localiza ions was la ge in males
(28
±
12 ha) han in emales (24 ±10 ha) du ing he whole o he
s udy pe iod, al hough he di e ence was no signi ican
(Table 2).
Homes anges and looding e ec s
The
la ges
home ange obse ed (48 ha) was ha o a emale
in he d y pe iod, which con as s wi h he la ges obse ed
male home ange (36 ha). Du ing he we season,
emales
also
had la ge home anges, wi h
a e ages
o 20 ha as opposed o
a e age male alues o 16 ha (Table 2). The di e ences in
Table 2 Home anges o adul Ibe ian ha es in eco one in sou he n Spain in wo seasons: d y (summe ), we (win e ) and home ange o al
Males
Females
N Min. Max. x
S
N Min. Max. x S p
MPC90 Eco one d y 5 12 36.2
22
8.9
9 13 47.6
22
12 ns
MPC90 Eco one we
5
10
23.5
16
5.6
3
14
25.4
20
5.5
ns
MPC90 Eco one To al
5
12
40.9
28
12
9
13
41
24
10
ns
Min minimum, Max m, x a e age, S s anda d de ia ion, MPC90 d y minimum con ex polygon wi h 90% o adio localisa ions in d y season,
MCP90 we minimum con ex polygon wi h 90% o adio localisa ions in we season, MPCTo al minimum con ex polygon wi h 90% o all adio
localisa ions, p signi ica ion, ns no signi ican
home anges obse ed in di e en seasons o he yea we e
also non-signi ican (Mann–Whi ney U es ,
U
=43;
p
=0.5).
Habi a use and habi a selec ion
Signi ican di e ences be ween sexes we e obse ed in he
use o sc ub a eas (Mann–Whi ney U
es
=
24;
p
=
0.016)
and in d y pas u es (Mann–Whi ney U
es
=
19;
p
=
0.004).
In bo h seasons, males mos o en used he pas u es o he
eco one and hen he ushes and sc ub. Males only used he
d y pas u es du ing he d y season, and no use o he s ands
o b acken o sedges was obse ed (Table 3).
O e all, sc ubland a eas we e used mo e du ing he we
season by males (Mann–Whi ney U
es
=
4.5;
p
=
0.08). No
signi ican di e ences we e obse ed be ween he d y and
we seasons o he han in he sc ub a eas, bo h sexes
selec ed ushes posi i ely
(I le
’
s
Index: +0.32 o males
du ing he d y season and +0.15 o emales in he we
season). O he habi a s we e ac i ely a oided (nega i e
I le
’
s
Index). S ands o b acken and sedges we e ne e
used by ei he sex. Sc ub was also la gely a oided, bu a
small pa was some imes used by males (Table 3).
Du ing he we season, ha es mo ed o highe a eas o
a oid loods and e en occupied a eas o sc ubland, whe eas
du ing he d y season ha es mo ed back owa ds he
ma shland and occupied a eas and plan communi ies ( o
example, eed beds) ha we e looded when wa e le els a e
high (Fig. 2). Ha es hus mig a ed in he d y pe iod owa ds
a eas ha became looded du ing he we season and in an
opposi e di ec ion du ing he we season (Table 4).
Discussion
The ha es sex and home ange size
We expec ed o ind signi ican a ia ion in home ange sizes
be ween sexes, seasons and habi a s. These
expec a ions
we e
Table 3 Habi a use o adul
Ibe ian ha e in he eco one (Ha)
du ing he we and d y seasons
N numbe o animals s udied;
Min minimum su ace a ea used
o co esponding plan commu-
ni y; Max minimum su ace a ea
used o co esponding plan
communi y;
A e age
a e age
su ace a ea used o
co esponding plan communi y;
x a e age; S s anda d de ia ion;
p signi icance le el; ns no
signi ican
We season D y season
N Min Max x S N Min Max x S p
Males
Sc ub 5 0 3.7 1.8 1.4 5 0 1.3 0.3 0.6 ns
Rushes 5 0.5 4.8 3.2 2.1 5 0 10.3 4.4 4.1 ns
D y pas u es 5 0 0 0 0 5 0 2 0.5 1.0 ns
B acken 5 0 0 0 0 5 0 0 0 0 ns
Sedges 5 0 0 0 0 5 0 0 0 0 ns
Reeds 5 0 0 0 0 5 0 0 0 0 ns
Pas u e in La Ve a 5 4.5 18.6 11.2 6.1 5 10.8 23.7 16.4 4.8 ns
Females
Sc ub 3 0 0 0 0 8 0 0 0 0 ns
Rushes 3 2.5 3.9 3.1 0.7 8 0.5 7.8 3.4 3 ns
D y pas u es 3 1 2 1.6 0.3 8 0 6 1.4 1.8 ns
B acken 3 0 0 0 0 8 0 0 0 0 ns
Sedges 3 0 0 0 0 8 0 0 0 0 ns
Reeds 3 0 0 0 0 8 0 1 0.1 0.3 ns
Pas u e in La Ve a 3 10.4 21.2 15.6 5.4 8 6.5 39 15.7 9.7 ns
p e e ence o habi a in he
Sc ub
Rushes
D y pas u e
B acken
Sedges
Reeds
Pas u e in eco one
Ibe ian Ha e in he eco one
zone o Doñana Na ional Pa k
Male WS
−0.96
0.17
−
1
−
1
−
1
−
1
−0.69
Male DS
−0.99
0.32
−0.56
−
1
−
1
−
1
−0.57
Female WS
−
1
0.15
−0.05
−
1
−
1
−
1
−0.59
Fig. 2 Habi a use in he d y season and in he we season in he eco one o pas u es be ween sc ubland (on he le ) and ma shland (on he igh
side o he igu e). Coo dina es UTM. Da um Eu opean 1959 ( o Spain and Po ugal)
based on p e ious published s udies ha epo ed a ma ked
in e - and in aspeci ic a ia ion in he size o home anges,
which may be explained by ac o s such as body weigh
(Ha es ad and Bunnell 1979), die and speci ic ene gy
equi emen s (McNab 1986).
P e ious in o ma ion on he home anges o he Ibe ian
Ha e is e y pa chy. Two unpublished epo s on ha es
eleased o
e-in oduc ion,
ci ed by Rod íguez e al. (1997),
epo ed home anges much la ge han hose ound in his
s udy (Fig. 3). Howe e , a compa ison be ween ou da a and
hose published by Rod íguez e al. (1997) is p oblema ical,
since hese au ho s wo ked wi h ha es whose beha iou may
ha e been a ec ed by ecen ansloca ion, likely causing
hem o pe o m e a ic mo emen s. Fu he mo e, home
ange size is ela ed o esou ce
a ailabili y
in la ge p eda o s
ein oduced
o Addo Elephan Na ional Pa k (Haywa d e al.
2009). No o he da a a e a ailable o L.
g ana
ensis.
Ibe ian ha e home ange s. o he ha es species
Compa able published da a om o he ha e species (Lepus
sp.) show ma ked a ia ion in home ange (Fig. 3), which is
acco ding o expec ed alues (Ha es ad and Bunnell 1979).
Adul Eu opean b own ha es (Lepus eu opaeus) a e 25–
35% la ge han adul Moun ain ha es (Lepus imidus) and
so will equi e la ge home anges. Ne e heless, hese
alues may a y depending on esou ces a ailable in he
habi a s in which he ha es li e. Eu opean ha es ha li e in
lowe -lying a eas wi h be e quali y ophic esou ces and a
g ea e a ie y o c oplands and nea by shel e a eas ha e
smalle home anges han hose ha li e in poo e -quali y
habi a s (Tappe and Ba nes 1986). Thus, in mosaic a eas
wi h an abundance o sh ubs, pas u es and c oplands, ha es
can shel e in a eas o co e and eed in nea by open a eas
wi hou ha ing o a el g ea dis ances.
Table 4
I le
’
s
Index o
Female DS −1 0.20 −0.12 −1 −1 −0.72 −0.59
WS
we
season,
DS
d y
season

Home ange (Ha)
Fig. 3 Home anges and mean
body size o ha e species as
epo ed in li e a u e. Pa kes
(1984); Pielowski (1972); Rei z
and Leona d (1994); Ma bou in
and Aebische (1996); Tappe
and Ba nes (1986); Ko acs and
Búza (1988); B oekhuizen and
Maaskamp (1982); Hulbe e al.
(1996); Hewson and Hinge
(1990);
O
’
Fa ell (1965);
Bou in (1984); Ca o (2005);
Ba is a and Mexia de Almeida
(1996), Fa án e al. (1999) and
Vi gós e al. (2006). Lam Lepus
ame icanus; Lg Lepus
g ana ensis; L i Lepus imidus;
160
140
120
100
80
60
40
20
0
Lam
Lg _ ansloca ed
L i
Lg _ his s udy
Leu
Leu Lepus eu opaeus 0 500 1000 1500 2000 2500 3000 3500 4000
4500
Weigh
(g )
Snowshoe ha es L. ame icanus need la ge home anges
gi en hei eliance as ood sou ces on hea he s, which
g ow in pa ches and may e en be le inaccessible by snow
co e (Hewson 1989). In a eas o abundan ophic
esou ces, hese ha es may e en en e in o compe i ion
wi h ca le (Hewson 1989). They also need shel e a eas
and will a el la ge dis ances o ind hem, mo ing
be ween a a ie y o eeding habi a s and e uge zones ha
p o ide shel e om he wind and o he me eo ological
e en s (Hewson and Hinge 1990).
Moun ain and Eu opean ha e emales ha e smalle home
anges han males. Hewson and Hinge (1990) ound ha in
hese species, home anges we e la ge du ing he b eeding
season, due o he ac , by an in e - and in a-sexual social
dominance hie a chy. Male home anges o e lap and males
sha e habi a (Hewson and Hinge 1990). In Doñana, we
ound no signi ican di e ences be ween home anges,
al hough emale home anges did end o be la ge . As
epo ed o o he species o ha es (Hewson 1986), male
and emale home anges coincide (bo h eeding and shel e
a eas) in bo h he d y and we seasons, and as such, no
e idence o e i o ial beha iou exis s (Fig. 2). Ne e he-
less, some con lic s be ween males ha lead o a hie a chy
in access o emales almos ce ainly occu .
In DNP, male and emale adul Ibe ian ha es di e in
habi a use. Ou da a showed ha males occasionally used
he co e and o he esou ces o sc ubland a ea while he
emales ne e we e obse ed ou side o he pas u e band.
Calzada E de la and Ma ínez (1994) s udied Ibe ian ha es
inhabi ing ano he ype o eco one using oad censuses.
They ound ha ha es ha e a hei disposal all he
en i onmen s hey need o be able o ca y ou hei li e
cycles and so ha e smalle home anges han in he la ge
ce eal plains o he Ibe ian Peninsula. These au ho s also
epo ed ha in cul i a ed a eas, home anges espond o he
cyclical changes ha occu in hese en i onmen s and
popula ions a e
es uc u ed
as hey mo e om un a ou able
o mo e a ou able li ing qua e s (de la Calzada and
Ma ínez 1994).
Rod íguez e al. (1997) eco ded e y la ge home anges
ha included a ious habi a ypes, which we e chosen
depending on equi emen s, al hough, as men ioned abo e,
he use o ansloca ed ha es makes he in e p e a ion o
hei da a complex. Home ange size depends on he ypes
and deg ee o homogenei y o he habi a (Ma bou in and
Aebische 1996) and he ime o ac i i y (Rei z and
Léona d 1994). Ou da a ag ee wi h he indings o Bou in
(1984) o L. ame icanus which was educe he home ange
size depending he op imal die . Ma shlands a e seconda y
habi a pa ially colonised e y d y season and holding high
densi y a e long d ough pe iods (Ca o 2005).
Ibe ian ha e home ange in we and d y season and looding
e ec
The a ia ions in home ange sizes be ween he d y and we
seasons we e no signi ican , al hough a ia ions did end o
be g ea es in he d y season, when pa o he eco one
loods in he we season, he amoun o a ailable o he
species is educed and ha es a e o ced o mo e o highe
and d ie a eas (Table 2). Wol (1980) ound ha L.
ame icanus occupied a eas ha p o ided e uge ( hick
o es ) in win e and mo e open a eas in summe , which
enabled ha es o espond o changes in he en i onmen .
The mac opodid ma supial quokka Se onix
b
a
ch
y
u u
s
inhabi s swamplands and exhibi s a simila mo emen
pa e n du ing seasonal looding (Haywa d e al. 2004).
Du ing ainy yea s, he e is high looding le el o he
ma shland ha ew a eas can be occupied by Ibe ian ha es.
When ha happens, Ibe ian ha es mo e (i hey can) om
inside
o he
ma shland
o each he
ma shland bo de ,
de ined
by eco one pas u es. These pas u es ac as a e uge habi a o
Ibe ian ha es in he a ea. I hey canno escape om he lood,
indi idual Ibe ian
ha es may ind also a
e uge
in he island o
ma shland, sca e ed
sligh ly highe
ele a ions
anging in size
om some dozens o me es o a ew hec a es, whe e hey
emain isola ed o mon hs, su e ing high
mo ali y
a es due
o p eda ion (Ca o e al. 2001, 2002; Ca o 2005). Only a
ew o hem, i any, su i e he long isola ion pe iod.
P eda ion and s a a ions a e he main mo ali y keys, same
o quokkas (Haywa d e al. 2005).
Acknowledgemen s The au ho s wish o exp ess hei g a i ude o
he s a —especially Jose J. Cháns and J.C Calde ón—o he Doñana
Na ional Pa k and Doñana Biological S a ion o hei help wi h he
ieldwo k. This s udy was pa ially suppo ed by he Plan Andaluz de
In es igación (Jun a de Andalucía; RNM-118) and P ojec
“
E ec
o
he loods on popula ions o ha es in he Doñana ma shland” CICYT
(1996
–
1997).
D Ma Haywa d and wo anonymous e e ees p o ided
insigh ul commen s on e sions o his manusc ip .
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