Sett Use, Density and Breeding Phenology of Badgers in Mediterranean Agro-Sylvo-Pastoral Systems

animals A icle Se Use, Densi y and B eeding Phenology o Badge s in Medi e anean Ag o-Syl o-Pas o al Sys ems Ma celo Sil a †, Luís Miguel Rosalino †, Sand a Alcobia and Ma ga ida San os-Reis *   Ci a ion: Sil a, M.; Rosalino, L.M.; Alcobia, S.; San os-Reis, M. Se Use, Densi y and B eeding Phenology o Badge s in Medi e anean Ag o-Syl o-Pas o al Sys ems. Animals 2021,11, 2663. h ps:// doi.o g/10.3390/ani11092663 Academic Edi o s: Nicola Ma ples, Aoibheann Gaugh an and Enda Mullen Recei ed: 1 Augus 2021 Accep ed: 9 Sep embe 2021 Published: 10 Sep embe 2021 Publishe ’s No e: MDPI s ays neu al wi h ega d o ju isdic ional claims in published maps and ins i u ional a il- ia ions. Copy igh : © 2021 by he au ho s. Licensee MDPI, Basel, Swi ze land. This a icle is an open access a icle dis ibu ed unde he e ms and condi ions o he C ea i e Commons A ibu ion (CC BY) license (h ps:// c ea i ecommons.o g/licenses/by/ 4.0/). cE3c—Cen e o Ecology, E olu ion and En i onmen al Changes, Faculdade de Ciências, Uni e sidade de Lisboa, 1749-016 Lisboa, Po ugal; [email p o ec ed] (M.S.); lm [email p o ec ed] (L.M.R.); [email p o ec ed] (S.A.) *Co espondence: mm [email p o ec ed] † These au ho s con ibu ed equally o his wo k. Simple Summa y: Unde s anding ca ni o es social s uc u e a ia ion is pi o al o p ope ly ad- d essing conse a ion challenges and solu ions. The Eu opean badge s is a social ca ni o e o which mos o he a ailable in o ma ion ega ding how his species is socially o ganized de i es om cen al wes popula ions. This a icle desc ibes he g oup composi ion, den use pa e ns and b eeding phenology o a Medi e anean popula ion o badge s. We showed ha badge li e in low densi y, in ela i ely small g oups, composed by 2–4 adul animals and ca. 2 cubs, bo n in win e . These pa e ns, ep esen ing a a ia ion o wha was desc ibed o o he popula ions, show ha badge s ake ad an age o he landscape con ex , whe e human- ela ed esou ces and mild en i onmen al condi ions allow badge o each highe densi ies han in many sou he n popula ions, and o ep oduce ea lie han hei no he n coun e pa s. Abs ac : Ca ni o es social o ganiza ion a ies widely, om s ongly social o soli a y p eda o s. Eu opean badge s a e acul a i e social ca ni o es ha also shows a geog aphical a ia ion in social s uc u e. These pa e ns de i e mainly om cen al/wes Eu opean egions, wi h an unde - ep esen a ion o Medi e anean popula ions ha ace di e en conse a ion challenges, especially ega ding g oup composi ion, se use pa e ns and b eeding phenology. We add essed hese ai s opics o a popula ion inhabi ing a Po uguese ag o-sil o-pas o al sys em. Based on mon hly moni o ing o 34 se s and con inuous came a- apping su eys o 12, we showed ha se s su - ounded by di e si ied ege a ion and loca ed in sandy si es a e mo e used, a pa e n p obably linked o ood a ailabili y and ease o se exca a ion and main enance, espec i ely. Badge s ol- lowed a gene al pa e n ega ding g oup size (2–4 adul s), bu showed an in e media e popula ion densi y ( 0.49–0.73 badge s/km2 ), wi h alues highe han hose es ima ed o o he Medi e anean en i onmen s, bu lowe han o cen al-wes e n popula ions. This, oge he wi h he b eeding (No embe /Janua y) and cub eme gence (1.8 cubs/se ; Ma ch/Ap il) pe iods, indica es an ecolog- ical adap a ion o he landscape con ex , whe e human- ela ed esou ces and mild en i onmen al condi ions allow badge o each highe densi ies han in many sou he n popula ions, and o ep o- duce ea lie han hei no he n coun e pa s. Keywo ds: came a- apping; Meles meles; densi y; social o ganiza ion; ep oduc ion 1. In oduc ion Ca ni o es show a wide a ay o social o ganiza ions, anging om soli a y (e.g., wild- ca s, Felis sil es is) o highly social species, such as he mee ka (Su ica a su ica a) [ 1 ]. Howe e , less han 20% a e conside ed social p eda o s [ 2 ], and e en hose show dis inc deg ees o sociali y. While wol es (Canis lupus) can o m packs wi h mo e han 40 animals sha ing hun ing, e i o y pa olling and de ense, cub ea ing, and o he ac i i ies [ 3 ], o he s, such as Eu opean badge s (Meles meles), o m clans usually wi h less han se en Animals 2021,11, 2663. h ps://doi.o g/10.3390/ani11092663 h ps://www.mdpi.com/jou nal/animals Animals 2021,11, 2663 2 o 15 indi iduals ha sha e he same e uge and e i o y and show some in a-clan in e ac ions, bu lack he social sophis ica ion showed by highly social species [ 4 ]. The way badge social g oups a e o ganized and use he communal e uges hey build (known as se s) is howe e highly a iable ac oss i s dis ibu ion ange and his a iabili y s ill poses ques ions abou how he landscape con ex may shape his species’ social pa e ns. Th oughou Eu ope, he size o badge social g oups anges om singe pai s in sou he n Spain [ 5 ] o se en adul indi iduals in Wy ham Woods, UK (al hough eaching 30 adul s/yea ling badge s in unusual si ua ions, such as hose ound in Woodches e Pa k, UK, in 1989) [ 6 ]. Such social s uc u e de e mines he species densi y, which peaks in he UK (38 ind./km 2 ) and eaches i s lowes alue in Eas e n Eu ope (e.g., Czech Republic, 0.12 ind./km 2 [ 7 ]. This a ia ion in densi y is equen ly linked o clima e a ia ion (e.g., we e clima e a o s ea hwo m’s abundance and, indi ec ly, badge s), abundance and a ailabili y o po en ial se si es and/o ood, and le el o dis u bance, such as human popula ion densi y, oad densi y o hun ing p essu e [8]. Ibe ia, co esponding o he species sou h-wes e n ange limi , seems o be a chal- lenging en i onmen o badge s, which occu a low densi ies (0.13–0.67 ind./km 2 , in Spain, and 0.36–0.48 ind./km 2 in Po ugal) [ 9 – 12 ]. This popula ion s uc u e seems o be de e mined mos ly by he low a ailabili y o ood esou ces [ 11 ] o se si es [ 13 ] in Medi e anean en i onmen s. Howe e , es ima ions o badge densi y a e limi ed o e y ew egions in Ibe ia—Doñana Na ional Pa k (S) and Pa k o Collse ola (NE), in Spain, and Se a de G ândola in Po ugal [ 10 – 12 ]—no ep esen ing he ull ange o en i on- men al condi ions badge s explo e in Ibe ia. Thus, mo e da a a e needed o allow a be e unde s anding o how Ibe ian popula ions espond o and a e in luenced by he Ibe ian landscape con ex , because his egion is expec ed o su e d as ic en i onmen al changes in he nex u u e due o clima e wa ming [14,15]. Se s play a cen al ole in badge socio-ecology, ac ing no only as e uge si es (whe e indi iduals spend as much as 70% o hei ime), bu also as ocal poin s o ep oduc ion, a nu se y, and a social exhibi ion and in e ac ion a ena [ 4 ]. Fo all hese easons, badge s densi y is o en linked o se densi y, especially when conside ing he main se s, i.e., la ge se s (>5 en ances) wi h signs o a egula use h oughou he yea [ 16 ]. Howe e , in low densi y a eas, as Ibe ia, his ela ion is weake , and keeps alid only including also smalle and less equen ly used se s (o en called, seconda y, ou lie s o subsidia y) [ 16 ]. Wha e e he ype o se s conside ed, hei use pa e n is de e mined by hei in e nal s uc u e, landscape con ex and dis u bance isk [17–19]. In he Medi e anean egion, human p esence has shaped he landscape o millennia, con e ing na u al en i onmen s o ag icul u e land, pas u e o li es ock o human se lemen s/in as uc u es [ 20 ], a ec ing wildli e ecological pa e ns. In Ibe ia, badge s show low se use a es, wi h animals equen ly changing be ween se s in consecu i e days (ca.50% o he occasions), and he e-use a e g ea ly declining a e i e nigh s [ 21 ]. This pa e n seems o be linked o an h opic dis u bances, such as ca le b eeding and emo al o o es unde s o y a ound he se s [22]. Se s a e also a pi o al s uc u e in badge ep oduc ion, as bi h occu s in unde - g ound se chambe s, and cubs s ay inside un il hey can explo e he ou side en i on- men [ 4 ]. While ma ing can occu h oughou he yea , mos a ailable bi h eco ds a e concen a ed in Feb ua y, due o delayed implan a ion, wi h a li e size o 2–3 cubs ( ange 1–5) [ 4 ]. Howe e , his in o ma ion o igina es mainly om UK popula ions, wi h sca ce da a a ailable o Medi e anean popula ions, whe e he highly di e en clima e may p oduce a ia ions o he B i ish pa e n. A ailable da a o he Ibe ian Peninsula indica e ha in cap i e animals ma ing occu ed be ween Feb ua y and May, wi h he bi h o 1–3 cubs in Feb ua y [ 23 ], while 3–4 cubs pe se ha e been epo ed o a wild popula ion in Po ugal [12]. Conside ing he speci ic cha ac e is ics o Medi e anean landscapes, and he lack o in o ma ion on he ecology o badge s in his egion, ou s udy aimed o assess badge : Animals 2021,11, 2663 3 o 15 (1) densi y; (2) se use pa e ns and unde lying de e minan s; and (3) b eeding phenology and li e size. To ul ill hese goals, we es ed eigh hypo heses (Table 1). Table 1. Hypo heses es ed a ge ing h ee li e-his o y pa e ns: se use, densi y and b eeding phenology. Fo each, we de ail he unde lying easoning and he suppo ing e e ences. Pa e n Hypo hesis Reasoning Suppo ing Re e ences Densi y H1—Badge ’s densi y will be lowe when compa ed o he species co e ange a ea, bu wi hin he limi s epo ed o Ibe ian popula ions S udies implemen ed in Ibe ia es ima e a popula ion densi y o 0.13–0.67 ind./km2, signi ican ly lowe han ha es ima ed o cen al/wes Eu opean popula ions (mean = 9.2 ind./km2, SD = 10.5) [4,9–12] Se use pa e n H2—Se use is p omo ed by he occu ence o dense ege a ion su ounding he se In human-shaped landscapes, he occu ence o dense ege a ion p o ides a mo e p o ec i e con ex , whe e animals may socially in e ac mo e sa ely (including cubs) [24,25] H3—Se s loca ed in a eas wi h a easily diggable and well-d ained soil will be mo e equen ly used Badge s p e e well d ained and cohesi e soils as se si es [26,27] H4—Se use is p omo ed by he p oximi y o eeding pa ches (e.g., oli e ya ds, wildli e eeding s a ions) Se loca ion nea pa ches p o iding ood esou ces allows badge s o sa e ene gy and ime in hei o aging bou s [22,28] H5—Se s subjec o high dis u bance (e.g., ca le, oads p oximi y) will be less used by badge s An h opogenic dis u bance is known o a ec se use by badge s [22,29–31] H6—Se s loca ed in a eas whe e pe cei ed compe i ion can be high will be less used E idence exis ha he p esence o wild boa s cons ains badge s p esence, due o esou ce compe i ion. In e -g oup compe i ion is also conside ed a passi e ange exclusion mechanism. [32,33] Rep oduc ion phenology H7—Badge ma ing peaks in Janua y/Feb ua y and cubs will s a eme ging om he se s in Ap il Badge ma ing can occu all yea ound, wi h wo peaks in Win e /Sp ing (main) and Summe /Au umn; cubs eme ge eigh weeks a e . [11,34] H8 —Li e s will be composed o 2–3 cubs Al hough a ailable da a o Po ugal indica e 3-4 cubs/li e , he a e age alues o mos badge popula ions is lowe [12,34] 2. Ma e ials and Me hods 2.1. S udy A ea The s udy was conduc ed in he “Cha neca do In an ado” (38 ◦ 48 0 N 8 ◦ 49 0 W; Figu e 1 ), a 100 km 2 a ms ead managed by Companhia das Lezí ias S.A., mainly de o ed o he p oduc ion o co k (co k oak, Que cus sube , woodlands—67.50 km 2 ), wood (ma i ime pine, Pinus pinas e , plan a ions—9.71 km 2 ; Eucalyp us, Eucalyp us globulus, plan a ions; 4.76 km 2 ), pine nu s (s one pine, Pinus pinea, plan a ions—5.08 km 2 ), and ag icul u al goods (e.g., co n, Zea mays—2.50 km 2 ; ice, O yza spp.—2.40 km 2 ; oli es, Olea eu opaea—0.70 km 2 ). This mul iuse ag o o es y sys em also includes o he ac i i ies, namely seasonal ca le aising, wi h ca le g azing in he a ms ead in Au umn/Win e , and hun ing mos ly o wild boa s (Sus sc o a) and pa idges (Alec o ys u a), o which se e al a i icial eeding s a ions wi h ce eals and wa e a e a ailable h oughou he a ms ead [35]. The egion has a ypical Medi e anean clima e, wi h ho and d y summe s and mild bu ainy win e s. Du ing he s udy pe iod he mean empe a u e and p ecipi a ion we e o 16.8 ◦ C and 361 mm, espec i ely. “Cha neca do In an ado” is loca ed in a plain lowland a ea, wi h ew ela i ely deep alleys o med by empo a y wa e cou ses. Only Animals 2021,11, 2663 4 o 15 one pe manen s eam c osses he a ms ead (“Ribei a de Vale Cob ão”), al hough se e al a i icial wa e poin s and empo a y ponds a e sca e ed h oughou he a eas. Animals2021,11,26634o 15  a i icial eedings a ionswi hce ealsandwa e a ea ailable h oughou  he a ms ead [35]. The egionhasa ypicalMedi e aneanclima e,wi hho andd ysumme sandmild bu  ainywin e s.Du ing hes udype iod hemean empe a u eandp ecipi a ionwe e o 16.8°Cand361mm, espec i ely.“Cha necadoIn an ado”isloca edinaplainlowland a ea,wi h ew ela i elydeep alleys o medby empo a ywa e cou ses.Onlyone pe manen s eamc osses he a ms ead(“Ribei adeValeCob ão”),al houghse e al a i icialwa e poin sand empo a ypondsa esca e ed h oughou  hea eas.  Figu e1.S udya ealoca ion(A),andspa ialdis ibu iono de ec edbadge se s(B),highligh ing hoseselec ed o  in ensi emoni o ingusingcame a‐ aps(C).Di e en colo s ep esen dis inc badge socialg oups,andmainse sa e ep esen edbybigge ci cles. 2.2.Se Su ey We i s  e iewedandcompiledallse  eco ds egis e edinp e iouss udies conduc edin hes udya ea(e.g.,[22,36]).All hesese s(n=33)we e e isi ed ocon i m hei usebybadge andnewse swe ein ensi elysea chedin hes udya ea.Thea eais di idedinca leg azingplo sand,be weenSep embe 2016andAp il2017,wesu eyed 20 ha  ul illed hecha ac e is icsconside edp omo e so se emplacemen ( ege a ion andsoil ea u es;see[4];anda oidingin ensi eag icul u ea eas,suchas iceandco n ields,wa e channeland ese oi s,e c.).Theseplo s, o alizing15km2(meana ea1.45 km2,sd=1.05, ange:0.05–6.66km2)we esu eyedbasedonse e allinea andpa allel ansec s,co e ing heen i ecell.Thedis ancebe ween he ansec swasabou 50me e s, Figu e 1. S udy a ea loca ion ( A ), and spa ial dis ibu ion o de ec ed badge se s ( B ), highligh ing hose selec ed o in ensi e moni o ing using came a- aps ( C ). Di e en colo s ep esen dis inc badge social g oups, and main se s a e ep esen ed by bigge ci cles. 2.2. Se Su ey We i s e iewed and compiled all se eco ds egis e ed in p e ious s udies conduc ed in he s udy a ea (e.g., [ 22 , 36 ]). All hese se s (n= 33) we e e isi ed o con i m hei use by badge and new se s we e in ensi ely sea ched in he s udy a ea. The a ea is di ided in ca le g azing plo s and, be ween Sep embe 2016 and Ap il 2017, we su eyed 20 ha ul illed he cha ac e is ics conside ed p omo e s o se emplacemen ( ege a ion and soil ea u es; see [ 4 ]; and a oiding in ensi e ag icul u e a eas, such as ice and co n ields, wa e channel and ese oi s, e c.). These plo s, o alizing 15 km 2 (mean a ea 1.45 km 2 , sd = 1.05 , ange: 0.05–6.66 km 2 ) we e su eyed based on se e al linea and pa allel ansec s, co e ing he en i e cell. The dis ance be ween he ansec s was abou 50 me e s, since we su eyed a bu e o 25 m in each pe pendicula di ec ion. In mo e closed a eas (woodlands, o example), he dis ance be ween ansec s was sho ened o ensu e ha he whole a ea was su eyed, as he isual de ec ion o se s dec eases in dense ege a ion a eas. Assuming ha main se s a e loca ed in he co e a ea o each g oup e i o y [ 24 ], and conside ing ha badge s home ange in a simila Po uguese landscape was 4.46 km 2 [ 12 ], Animals 2021,11, 2663 5 o 15 we assumed ha all seconda y se s loca ed wi hin a adius o 1191 m om a main se (see Sec ion 2.3) will belong o he same social g oup. The only excep ions we e seconda y se s ei he sepa a ed by an insu moun able geog aphical ba ie (e.g., pe manen s eam) o loca ed wi hin he bu e bounda ies o wo neighbo main se s (in such si ua ion hey we e g ouped wi h he closes main se ). 2.3. Se Moni o ing To de e mine he se use pa e n, all de ec ed se s we e isi ed mon hly o de e mine badge signs o p esence, and es ima e a se use index, anging om 0 (ne e ac i e) o 7 always ac i e. A se was conside ed ac i e i nea by i s en ances we ound: (1) eshly exca a ed soil, (2) ecen ly used la ines; (3) esh nes ma e ial (Figu e S1; Supplemen a y Ma e ial); and/o (4) high abundance o oo p in s. Se s ha showed equen and abundan ac i i y signs (se index = 6–7), o whe e e- p oduc ion was de ec ed, we e conside ed main se s [ 12 ], all he emaining being classi ied as seconda y [ 24 ]. The main se s we e he a ge o an in ense moni o ing p og am, using a came a apping app oach [ 37 ]. We ins alled 14 Moul ie ® m-990i Gen2 (Cale a, AL, USA) came a- aps (Figu e S2; Supplemen a y Ma e ial), equipped wi h a 16 Gb memo y ca d, se o ake 30s low esolu ion ideos (854–480 pixels) when he hea /mo emen senso was ac i a ed, wi h a minimum ime in e al be ween consecu i e ideos o 5s. Came as we e ins alled 30 cm abo e he g ound, a ached o a ee o a wooden s ick, acing one o mo e ac i e se en ances. In wo se s wi h mo e han one en ance ac i e, and ha could no be simul aneously moni o ed wi h a single came a, an addi ional one was se o ensu e he moni o ing o all ac i e en ances. Came as we e checked weekly o eplace he ba e ies and memo y ca ds. Se moni o ing occu ed be ween No embe 2016 and Ap il 2017, and each came a was ac i e o 180 days, o an o e all moni o ing pe iod o 2520 came a- apping nigh s. 2.4. Se Cha ac e iza ion In o de o assess which en i onmen al d i e s, we e explaining he se use pa e n, we used 21 a iables o cha ac e ize each se (Table 2). Va iables associa ed o ege a ion composi ion and s uc u e we e assessed, in si u, wi hin he minimum con ex polygon ha encompassed all he se en ances, plus a bu e o 2 m. The Simpson di e si y index [ 38 ] was es ima ed based on he he baceous, sh ub and ba e soil co e in each polygon. La e , we buil a Geog aphic In o ma ion Sys em (GIS; QGIS ® e sion 2.8.3—Wien, Bea e on, OR, USA [ 39 ]), ha encompassed emo e sensing in o ma ion ega ding he: ype o soil (da a p o ided by he Companhia das Lezí ia, S.A, Samo a Co eia, Po ugal); land use [ 38 ]; loca ion o se s, game eeding s a ions and wa e ing pi o s, pa ed and unpa ed oads and wa e poin s ( o allow he es ima ion o he dis ance o each se om hese landma ks; da a p o ided by he Companhia das Lezí ia, S.A.); g azing p essu e pe plo (da a p o ided by he Companhia das Lezí ia, S.A.); and wild boa occu ence de e mined in a p e ious s udy [40]. 2.5. G oup Size and B eeding Phenology Each ideo was ca e ully iewed and we egis e ed he numbe o indi iduals iden- i ied, he age class (i.e., adul , cub; based on animal’s size and su ey mon h [ 4 ], he in e ac ion be ween he animal and he se (i.e., en e ing o lea ing he se , exca a ing o handling nes ma e ial), as well o he beha io s o social in e ac ions (e.g., an agonis ic o b eeding beha io s). To es ima e he numbe o indi iduals pe social g oup we used he highes numbe o badge s eco ded simul aneously. Howe e , his es ima ion was only conside ed alid i ha numbe o indi iduals was obse ed in, a leas 1% o he o al eco ded ideos o each se . This c i e ion was de ined o educe he isk o coun ing, as membe s o he moni o ed g oup, indi iduals om o he social g oups ha may be isi ing he main se o ano he g oup. This conse a i e app oach conside s ha all badge s de ec ed simul aneously belong o he same social g oup. Animals 2021,11, 2663 6 o 15 Table 2. Va iables used o cha ac e ize each se du ing he mon hly moni o ing isi , wi h e e ence o he hypo hesis o which hey we e linked, he ype o a iable and i s ange/uni s. Va iable Desc ip ion Type Range/Uni s H2 T ee T ee co e O dinal 1–5 (1—low; 5—high) 1,2 Sh ubs Sh ub co e Con inuous Pe cen age 1,2 He bs He baceous co e Con inuous Pe cen age 1,2 Sh ub_H Sh ub mean high Con inuous Cm 1,2 Ba e_soil Ba e soil co e Con inuous Pe cen age 1,2 Veg_di Vege a ion di e si y Con inuous 0–1 (Simpson di e si y index) Unde s o y Dominan unde s o y Ca ego ical 4 Ca ego ies (Ca le pas u es, Sh ubland, Na u al pas u es, Na u al pas u es wi h sh ubs) Land use Type o land use Ca ego ical 4 ca ego ies (Co k oak woodland, Mixes wood, Na u al pas u e, Pine o es ) H3 Soil_ma Main soil ma e ial Ca ego ical 4 ca ego ies (Sand, Rock, Sil /Clay, Roads 3)1 Soil Type o soil 2 Ca ego ies (Podzols, Regosols) H4 Dis _ol Dis ance o he nea es oli e ya d Con inuous m Dis _pi Dis ance o he nea es wa e ing pi o Con inuous m Dis _ eed Dis ance o he nea es game a i icial eede s Con inuous m Dis _wa Dis ance o he nea es wa e sou ce Con inuous m H5 G azz_16–17 G azing p essu e be ween 2016–2017 Con inuous G azing in ensi y 4 G azz_07–17 Cumula i e g azing p essu e be ween 2007–2017 Con inuous G azing in ensi y 4 N_g azz Numbe o consecu i e yea s, since 2007, wi hou ca le g azing Con inuous Numbe o yea s Dis _ oad Dis ance o he nea es pa ed oad Con inuous m Dis _ oad2 Dis ance o he nea es di oad Con inuous m H6 Dis _se Dis ance o he nea es badge se Con inuous m Wildboa Wild boa abundance Con inuous Numbe o wild boa signs [40] 1 Va iables collec ed in si u, wi h he minimum con ex polygon encompassing all he se en ances. 2 Va iables used in he P incipal Componen s Analysis (PCA). 3 Se s loca ed unde di oads. 4 Exp essed as he numbe o ca le pe day and pe hec a e, in he po whe e each se was loca ed [35]. Animals 2021,11, 2663 7 o 15 Rega ding ep oduc ion, we conside ed ha a ma ing e en occu ed when we de- ec ed in he ideo a sexual in e ac ion be ween wo adul badge s. When in subsequen ideos he ma ing e en s ill occu ed, we egis e ed he en i e sequence as a single ma ing e en . Ma ing e en s less han 2 minu es long we e conside ed sho -du a ion ma ing o copula ion e en s, and hose ha occu ed o mo e han 2 minu es ( ypically 12 minu es o mo e; see esul s) we e egis e ed as long-du a ion ma ing e en s [ 34 ]. Long ma ing e en s a e expec ed o inc ease b eeding success [41]. 2.6. Da a Analysis 2.6.1. D i e s o Se Use In he modelling p ocedu e applied o assessing he en i onmen al d i e s o se use by badge s (hypo heses H2–H6), he numbe o mon hly isi s posi i e o signs o ac i i y ( anging om 0 o 7) was included as dependen a iable (Tables 1and 2). Due o he high numbe o a iables associa e o H2 we conduc ed a P incipal Compo- nen Analysis (PCA) [ 42 ], using he a iables ee, sh ub, he baceous and ba e soil co e and sh ub heigh (Table 2); p incipal componen s ha cumula i ely explained >80% o he in o ma ion o he o iginal a iables, we e used as candida e d i e s in he subsequen modelling p ocedu e (see 3.1). Spa ial au oco ela ion o da a was assessed using he Mo an I Index [ 43 ], and we e al- ua ed mul icollinea i y be ween all candida e d i e s (excep hose used in he PCA) using he Spea man co ela ion coe icien , ρ [ 42 ]. When wo a iables we e highly co ela ed (ρ> 0.70), we excluded he one less co ela ed wi h he dependen a iable [44]. Fo each hypo hesis, we un Gene alized Linea Models (GLM) [ 42 ], using a Poisson dis ibu ion and a “log” link unc ion, co esponding o all combina ion o he a iables associa ed o he hypo hesis (Table 2). Resul ing models we e anked acco ding o he Akaike In o ma ion C i e ion, wi h co ec ion o small sample size (AICc) [ 45 ]; hose ha showed a ∆ AICc < 2 (i.e., he di e ence be ween he AICc o a model and he lowes AICc alue in he model se [ 46 ]) we e conside ed he bes models in explaining se use a ia ion. I mo e han one model ul illed his c i e ion, we used a model a e aging p ocedu e [47] o assess a iables a e age coe icien and he 95% con iden in e al (95% CI). Va iables wi h a 95% CI ha did no include he 0 (i.e., we can de e mine mo e p ecisely i i s e ec is posi i e o nega i e), and ha showed a highe ela i e impo ance, we e conside ed mo e in luen ial [ 48 ]. Rela i e impo ance was es ima ed as he sum o he Akaike weigh (w; p obabili y o a model being he bes model) [ 45 ], o all he models ha included he a iable o in e es . The a iables ha , o each hypo hesis, showed a signi ican (p< 0.05) o almos signi ican (p< 0.1) in luence on se use pa e n, we e included in o e all models (combined hypo hesis), ha pos ula ed ha he pa e n o se use is de e mined by a combina ion o d i e s linked o dis inc ecological/en i onmen al p ocesses ( ege a ion, soil, ood, dis u bance o compe i ion). Finally, we compa ed he AICc o he bes model o each hypo hesis and conside ed he hypo hesis wi h mo e suppo as ha showing he lowes AICc alue. The goodness o i o he bes o e all model was assessed by es ima ing he R 2 , which iden i ies he p opo ion o a iabili y o he o iginal da a explained by he model [ 47 ]. Finally, we also es ed he au oco ela ion o models esiduals [46]. 2.6.2. Badge Densi y Es ima ion We used he maximum numbe o di e en animals de ec ed in each main se (see abo e) o es ima e he mean numbe o animals pe social g oup o he en i e s udy a ea. We hen es ima ed badge densi y as: Mean numbe o adul badge s pe main se es ima ed o he s udy a ea ×Es ima ed numbe o social g oups Su eyed a ea 43 km2 Animals 2021,11, 2663 8 o 15 We assumed ha he s udied badge popula ion ollowed he beha io al pa e n desc ibed o o he a eas o i s dis ibu ion ange, wi h each social g oup ha ing one main se , used all yea ound and whe e ep oduc ion occu s [49]. 3. Resul s O a o al o 33 se s iden i ied in he a ea in he ame o p e ious s udies, only 22 s ill occu ed (i.e., we e no des oyed). Twen y- h ee new se s we e added as a esul o he ield wo k, o alizing 45 se s o moni o ing pu poses (Figu e 1B), esul ing in a se densi y o 1.05 se s/km2. 3.1. D i e s o Se Use Pa e n F om he 45 se s we de ec ed in he s udy a ea, we only used 34 in ou modeling p o- cedu e, since 11 we e only de ec ed close o he end o he s udy no p o iding obus da a ega ding hei use by badge s. No signi ican spa ial au oco ela ion was de ec ed (Mo an I = − 0.053; p= 0.822). F om all he candida e co- a ia es we excluded “G azz_07–17” and “G azz”, since hey we e signi ican ly co ela ed wi h “G azz_16–17 ( G ass_07–17–G azz = 0.77, p< 0.001; G ass_07–17–G azz_16–17 = 0.77, p< 0.001; G azz–G azz_16–17 = 0.79 , p< 0.001), and he la e was mo e co ela ed wi h he dependen a iable ( s = 0.258). Fu he mo e, we also excluded “Dis _pi ” as i was signi ican ly co ela ed o “Dis _ol” ( = 0.97, p< 0.001), and he “Dis _ol” was mo e co ela ed o he equency o se use ( s = −0.237). The i s wo compomen s o PCA analysis p oduced o he ege a ion associa ed a iables eached a cumula i e explained a iance o 80.1% and hus we e used as candi- da e a iables in models p ocuced o es Hypo hesis 2 (Vege a ion d i e s). The associa ion be ween bo h componen s and he o iginal a iables is p esen ed in Table 3. Table 3. Co ela ion be ween he o iginal a iables and he PCA componen s used in he modeling p ocedu e. Va iable desc ip ion is de ailed in Table 2. O iginal Va iable Componen 1 Componen 2 T ee −0.431 - Sh ubs −0.501 −0.398 He bs −0.507 −0.132 Sh ub_H −0.547 −0.271 Ba e_soil −0.114 0.865 We p oduced a o al o 91 models, bu only a ew we e conside ed he bes models (i.e., ∆ AICc < 2) o each hypo hesis: ou o H2, one o H3, wo o H4, h ee o H5 and wo o H6 (Table 4). F om he a iables included in hese bes models, only “Vege a ion di e si y”, “Dis ance o he nea es oli e ya ds”, ”Main soil ma e ial”, “Dis ance o he nea es pa ed oad”, “Dis ance o he nea es di oad” and “G azing p essu e in 2016–2017 ” had a signi ican in luence in he dependen a iable (p< 0.001) and hus we e used as candida e a iables o he combined model (Table 4). Models o he combined hypo hesis showed he highes i (i.e., lowe AICc and O e all ∆ AICc < 2). Bes models included he a iables “Main soil ma e ial”, “Vege a ion di e si y”, “Dis ance o he nea es di oad” and “G azing p essu e in 2016–2017”. Based on he model a e aging p ocedu e we can s a e ha se s ha we e mo e used by bad- ge s du ing he s udy pe iod we e loca ed in si es wi h a highe di e si y o ege a ion (i.e., highe Simpson di e si y index), highe le els o g azing in he 2016–2017, and so e soils (e.g., sil /clay soils we e a oided) espec o poo ly used se s (Table 5). The bes model was able o explain ca. 40% o he a iance in he equency o se use (R 2 = 0.397), indica ing an accep able goodness o i [ 42 ], and no spa ial au oco ela ion o esiduals was de ec ed (Mo an I = −0.09; p= 0.55). Animals 2021,11, 2663 9 o 15 Table 4. Bes models o each es ed hypo hesis ( ∆ AICc < 2). Fo each model we include he deg ee o eedom (d ), he Akaike In o ma ion C i e ion, co ec ed o small samples (AICc), he AICc di e ence o he lowes AICc in each hypo hesis ( ∆ AICc), he Akaike weigh (Weigh ), and he AICc di e ence o he lowes AICc o all p oduced models (O e all ∆ AICc). Va iables’ ac onyms a e desc ibed in Table 2. Model d LogLik AICc ∆AICc Weigh O e all ∆AICc Null 1 −87.721 169.6 0 - 11.0 H2—Vege a ion Unde s o y + Veg_di + PCA1 6 −73.950 163.0 0 0.209 4.4 Unde s o y + Veg_di 5 −75.574 163.3 0.3 0.182 4.7 Veg_di 2 −70.981 164.3 1.3 0.107 5.7 Veg_di + PCA1 3 −78.977 164.8 1.8 0.088 6.2 H3—Soil Soil_ma 4 −75.027 159.4 0 0.781 0.8 H4—Food Dis _ol 2 −82.114 168.6 0 0.346 10.0 Dis _ol + Dis _wa 3 −81.902 170.6 2.0 0.128 12.0 H5—Dis u bance Dis _ oad2 + G azz_16–17 3 −78.930 164.7 0 0.255 6.1 Dis _ oad2 + G azz_16–17 + N_g azz 4−78.370 166.1 1.4 0.123 7.5 G azz_16–17 2 −81.036 166.5 1.8 0.104 7.9 H6—Compe i ion Dis _se 2 −83.464 171.3 0 0.223 12.7 Wildboa 2 −83.715 171.8 0.5 0.174 13.2 Combined hypo hesis Soil_ma + Veg_di 5 −73.204 158.6 0 0.151 0 Dis _ oad2 + G azz_16-17 + Veg_di 4 −74.792 159.0 0.4 0.123 0.4 Soil_ma + G azz_16-17 + Veg_di 6 −72.095 159.3 0.7 0.104 0.7 Soil_ma 4 −75.027 159.4 0.8 0.097 0.8 Veg_di + G azz_16-17 3 −76.340 159.5 0.9 0.095 0.9 Table 5. Va iables included in he a e age model o he bes models p oduced o explain he a ia ion in se use equency (Combined hypo hesis—Table 4). Fo each a iable he coe icien ( β ), s anda d e o (SE), z- alue and signi icance (p), 95% con idence in e als (95% CI) and ela i e impo ance a e p esen ed. (See Table 2 o a iables’ desc ip ion). Va iable βSE z-Value p95% CI Rela i e Impo ance In e cep 0.719 0.601 1.175 0.240 [−0.480, 1.919] - Soil_ma (Sil /clay) −0.740 0.244 2.919 0.004 [−1.237, −0.243] 0.62 Soil_ma (Rock) −0.115 0.246 0.450 0.653 [−0.618, 0.388] 0.62 Soil_ma (Roads) 0.264 0.403 0.629 0.530 [−0.559, 1.087] 0.62 Veg_di 1.989 0.862 2.122 0.034 [0.145, 3.651] 0.83 Dis _ oad2 −0.157 0.091 1.662 0.096 [−0.342, 0.028] 0.22 G azz_16-17 0.194 0.092 2.041 0.041 [0.007, 0.381] 0.56 3.2. Social G oups Size and Densi y Es ima es We iden i ied 12 se s ha showed a highe and mo e con inuous ac i i y pa e ns (Figu e 1C) and we e he e o e conside ed main se s. On a e age main se s included 3.85 ±3.32 ( anging om 1–7) en ances, 3 ± 2.2 o which we e ac i e simul aneously (i.e., wi h oo p in s, la ines, e ol ed soil, bedding ma e ial; ange 1–6). Seconda y se s had only an a e age o 1.41 ± 1.66 ac i e en ances, anging om 0 o 4 (o en none showed signs o ac i i y), om an o e all 2.93 ± 2.80 o al en ances ( ange: 1–9). The index o se use was highe o main se s (6.14 ±1.22) han o seconda y se s (3.37 ±2.23). O e all, we manage o egis e badge s ac i i ies in main se s on a e age 150 ±26 nigh pe se , ep esen ing 18,966 ideos o 30 seconds (i.e., 158 hou s o ideo). In 46% o he ideos (n= 8654) we de ec ed a leas one badge , and in 7% (n= 1281) o he mammal species: ca le, wild boa s, wild abbi (O yc olagus cuniculus), common gene (Gene a gene a), s one ma en (Ma es oina), ed ox (Vulpes ulpes) and Egyp ian mongoose (He -