Heavy flooding effects on home range and habitat selection of free-ranging Iberian hares (Lepus granatensis) in Doñana National Park (SW Spain)

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 . Re e ences Ba is a T, Mexia de Almeida L (1996) Resul ados das medidas e ou as obse açoes em leb es cob adas em p o as e caçadas, de 1992 a 1995. Re Flo es 9(1):123–135 Bou in S (1984) E ec o la e win e ood addi ion on numbe s and mo emen s o snowshoe ha es. 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