Patterns and Drivers of Rodent Abundance across a South African Multi-Use Landscape

animals A icle Pa e ns and D i e s o Roden Abundance ac oss a Sou h A ican Mul i-Use Landscape Bea iz C. A onso 1,* , Lou ens H. Swanepoel 2, Bea iz P. Rosa 1, Tiago A. Ma ques 3,4, Luís M. Rosalino 1, Ma ga ida San os-Reis 1and Gonçalo Cu ei a-San os 1   Ci a ion: A onso, B.C.; Swanepoel, L.H.; Rosa, B.P.; Ma ques, T.A.; Rosalino, L.M.; San os-Reis, M.; Cu ei a-San os, G. Pa e ns and D i e s o Roden Abundance ac oss a Sou h A ican Mul i-Use Landscape. Animals 2021,11, 2618. h ps:// doi.o g/10.3390/ani11092618 Academic Edi o : Emiliano Mo i Recei ed: 2 Augus 2021 Accep ed: 31 Augus 2021 Published: 7 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/). 1 cE3c—Cen e o Ecology, E olu ion and En i onmen al Changes, Faculdade de Ciências da Uni e sidade de Lisboa, Campo G ande, 1749-016 Lisboa, Po ugal; [email p o ec ed] (B.P.R.); [email p o ec ed] (L.M.R.); mm [email p o ec ed] (M.S.-R.); [email p o ec ed] (G.C.-S.) 2Depa men o Zoology, School o Ma hema ical & Na u al Sciences, Uni e si y o Venda, Thohoyandou 0950, Limpopo, Sou h A ica; [email p o ec ed] 3Cen e o Resea ch in o Ecological and En i onmen al Modelling, The Obse a o y, Uni e si y o S And ews, S And ews KY16 9LZ, UK; iago.ma [email p o ec ed] 4Cen o de Es a ís ica e Aplicações, Depa amen o de Biologia Animal, Faculdade de Ciências, Uni e sidade de Lisboa, 1749-016 Lisboa, Po ugal *Co espondence: bea izca [email p o ec ed] Simple Summa y: Wildli e ecological pa e ns a e d i en no only by en i onmen al and biological con ex s, bu also by landscape-managemen schemes ha shape hose con ex s. The p esen s udy aims o de e mine he e ec o di e en en i onmen al ac o s (including managemen schemes) on he occu ence pa e ns o a sou he n A ican small mammal communi y. Based on a landscape whe e h ee land-use con ex s ha di e in hei le els o human p esence and/o whe e ac i i ies coexis (p i a e eco ou ism ese e, mixed a ms and adi ional communal a eas), and by using a body-size-based app oach (i.e., using wo size-based oden g oups—medium and small—as models), we ound ha he mean ela i e abundance o medium-sized species did no di e ac oss he managemen con ex s, bu small species’ mean ela i e abundance was highe in he game ese e. The o e all a ia ion in oden abundance was nega i ely a ec ed by ungula e p esence (possibly linked o a dec ease in ood a ailabili y) and by human p esence (inc eased dis u bance). Roden abundance seems o be in luenced by en i onmen al g adien s ha a e di ec ly linked o a ying managemen p io i ies ac oss land uses, meaning ha hese communi ies migh no bene i uni o mly by he inc eased amoun o habi a p omo ed by he comme cial wildli e indus y. Abs ac : Sou h A ica’s decen alized app oach o conse a ion en ails ha wildli e ou side o mally p o ec ed a eas inhabi complex mul i-use landscapes, whe e p i a e wildli e business (eco ou ism and/o hun ing) co-exis in a human-domina ed landscape ma ix. Unde decen alized conse a ion, wildli e is pe cei ed o bene i om inc eased amoun o a ailable habi a , howe e i is c ucial o unde s and how dis inc managemen p io i ies and associa ed landscape modi ica ions impac noncha isma ic axa, such as small mammals. We conduc ed ex ensi e ink- acking- unnel su eys o es ima e he e ogenei y in oden dis ibu ion and in es iga e he e ec o di e en en i onmen al ac o s on abundance pa e ns o wo size-based oden g oups (small- and medium-sized species), ac oss h ee adjacen managemen con ex s in NE KwaZulu-Na al, Sou h A ica: a p i a e eco ou ism game ese e, mixed a ms and adi ional communal a eas (consis ing o small clus e s o houses in e spe sed wi h g azing a eas and semina u al ege a ion). Ou hypo heses we e o mula ed ega ding he (1) a ea ypology, (2) ege a ion s uc u e, (3) ungula e p essu e and (4) human dis u bance. Using a boos ed- eg ession- ee app oach, we ound conside able di e ences be ween oden g oups’ abundance and dis ibu ion, and he unde lying en i onmen al ac o s. The mean ela i e abundance o medium-sized species did no di e ac oss he h ee managemen con ex s, bu small species mean ela i e abundance was highe in he game ese es, con i ming an in luence o he a ea ypology on hei abundance. Va ia ion in oden ela i e abundance was nega i ely co ela ed wi h human dis u bance and ungula e p esence. Roden abundance seems o be in luenced by en i onmen al g adien s ha a e di ec ly linked o a ying managemen p io i ies ac oss land Animals 2021,11, 2618. h ps://doi.o g/10.3390/ani11092618 h ps://www.mdpi.com/jou nal/animals Animals 2021,11, 2618 2 o 18 uses, meaning ha hese communi ies migh no bene i uni o mly by he inc eased amoun o habi a p omo ed by he comme cial wildli e indus y. Keywo ds: non-in asi e sampling; ecological modelling; managemen op ions; conse a ion 1. In oduc ion In Sou h A ica, ag icul u al in ensi ica ion, and o e g azing ha e led o p o ound land use changes [ 1 ]. His o ically, mos landscapes we e con e ed in o li es ock a ms and a mlands, ei he as in ensi e, ex ensi e, o communally managed a eas [ 2 ], leading o he des uc ion, deg ada ion and/o agmen a ion o na u al ecosys ems [ 3 ]. Consequen ly, such habi a des uc ion led o declines in wildli e popula ions and dis ibu ion in much o Sou h A ican nonp o ec ed a eas [4]. Howe e , he es ablishmen o na ional policies a ibu ing cus odial igh s o e wildli e o landowne s, p omp ed a ansi ion in he go e nance o na u al esou ces om he s a e o p i a es [ 5 ]. This poli ical op ion led o widesp ead con e sion o angelands, i.e., a mlands and li es ock a ms, in o a eas dedica ed o comme cial wildli e indus ies, such as game anching and p i a e game/eco ou ism ese es [ 6 ]. The posi i e conse a- ion ou comes o hese policies o economically aluable and cha isma ic species [ 7 ] is belie ed o ha e an umb ella e ec on o he axa, mainly h ough he inc eased co e age, ep esen a i eness and connec i i y o p o ec ed/ es o ed habi a s [ 6 , 8 ]. Howe e , he e - ec o such managemen app oaches is unexplo ed o mos o e looked—bu unc ionally impo an — axa, such as oden s [ 9 ]. Thus, in o ma ion on he ecological esponses o less-cha isma ic axa is needed o be e gauge he complemen a y conse a ion ole o Sou h A ica’s p i a e land. In Sou h A ica, game a ms and p i a e game ese es o en coincide ac oss ela i ely small scales, oo ed in human-domina ed landscapes (e.g., communal lands) [ 10 ]. These land uses ha e con as ing managemen p io i ies and, consequen ly, dis inc impac s on he landscape s uc u e and wildli e ecological pa e ns. In game a ms, he main objec i e is o maximize he p oduc ion o ungula es o mea o hun ing, while in p i a e game ese es he goal is o main ain cha isma ic species, p omo ing eco ou ism-based ac i i- ies [ 11 ]. O en, hese wildli e-o ien ed land uses a e su ounded by human-domina ed a eas wi h high le els o an h opogenic dis u bance. The egional co-exis ence o all hese land uses gene a es complex mul i- enu ed landscapes, usually di ided by semi-pe meable wildli e ences, in luencing he biodi e si y suppo ed by each o hese land uses [12]. Managemen ac ions di ec ed o cha isma ic o aluable species may ha e cascading e ec s on oden s, usually o e looked and handled like pes s [ 13 – 15 ]. Howe e , i is c ucial o unde s and he e ec o human-induced land-use changes on oden spa ial pa e ns, as well as he unde lying ecological mechanisms he eo , since oden s a e undamen al o some ecosys em unc ions [ 16 ]. Roden s a e p ima y consume s [ 16 ] and suppo a la ge communi y o p eda o s [ 17 , 18 ], which makes hem a i al link in ood-chain s uc u ing [ 19 ]. Mo eo e , hey a e conside ed use ul indica o s o ecosys em unc ioning as hey a e aluable ools o he desc ip ion and moni o ing o habi a in eg i y. Fo hese easons, oden s ha e been used as model species o unde s and how land use changes a ec s wildli e [16]. Se e al ac o s ha e been iden i ied as in luen ial in shaping oden communi y and popula ion s uc u es, many o which a e o en de e mined by he landscape managemen op ions [ 20 ]. Some s udies ha e indica ed ha ege a ion ype and s uc u e a e undamen- al d i e s o oden occu ence and abundance [ 21 – 23 ]. Fo example, a eas wi h g ea e he baceous co e age a o oden s by p o iding shel e agains p eda o s, ood, and ade- qua e mic oclima ic condi ions [ 24 ]. S udies ha e shown nega i e e ec s o o e g azing on small mammals’ abundance, by educing he he baceous s a um, inc easing ampling isk and eeding compe i ion wi h ungula es [ 9 , 24 – 27 ]. Rega ding oden dis ibu ion, i ends Animals 2021,11, 2618 3 o 18 o be uni o m when he habi a is a o able and esou ces a e abundan . Howe e , when dis u bances inc ease he le el o habi a he e ogenei y, causing landscape agmen a ion, hei dis ibu ion is mos ly clumped [28,29]. Roden s a e no a homogeneous g oup, since di e en species may es ablish dis inc ela ionships wi h he en i onmen al and bio ic componen s o he ecosys em. Fo example, la ge oden s’ ange o e la ge spa ial scales han smalle oden s [ 30 ] and, he e o e, a e mo e suscep ible o changes a his landscape le el [31]. Changes in managemen p io i ies ac oss Sou h A ican mul i- enu ed landscapes will ha e a di ec impac on hese en i onmen al d i e s and, ul ima ely, in he dis ibu ion and abundance o small mammal species ac oss and wi hin managemen con ex s. Fo ins ance, when managemen measu es p omo e he abundance o ungula es (e.g., as p ey o la ge ca ni o e popula ions in eco ou ism ese es, o as hun ing asse s in game a ms), g azing p essu e will inc ease, nega i ely in luencing he he baceous s a a [ 26 ]. Alongside wi h long d y and ho seasons [ 32 ], hese condi ions may lead o sh ub enc oachmen , known o educe ood a ailabili y (lea es, seeds, and a h opods) o g ound dwelling oden s [ 33 ]. Ne e heless, some oden species a e usually conside ed e icien colonize s o human shaped en i onmen s [ 13 , 34 ], as hey a e able o use human- ela ed ood esou ces due o hei omni o e cha ac e [35]. Al hough he p ocesses ha egula e small mammals’ spa ial dis ibu ion a e known o some landscapes (e.g., woodland [ 29 ] and mixed o es [ 36 ]), he e is a lack o in o ma- ion ega ding he d i e s o oden -abundance pa e ns in A ican sa annas (bu see [ 9 , 37 ]), as well as how hese a y ac oss di e en managemen schemes. He e, we e alua ed he a ia ion in oden abundance ac oss h ee adjacen managemen con ex s, spanning a p i a e eco ou ism game ese e, mixed a ms and communally owned land, managed by Zulu ibal au ho i ies [ 12 ], unde he ollowing wo main objec i es: (1) o es ima e he e ogenei y in small-mammal-abundance dis ibu ion (mean abundance and pa chiness) ac oss managemen con ex s (game ese e, mixed a ms and communal lands); and (2) o de e mine he main, ine-scale en i onmen al ac o s a ec ing small-mammal-abundance pa e ns ac oss land-use ypes. These objec i es we e es ed in wo size-based oden g oups, o a mo e de ailed assessmen o ecological esponses. Linked o hese wo goals, we es ed ou hypo he ical d i e s o oden communi ies: (i) An a ea- ypology hypo hesis, i.e., cumula i e e ec o managemen -induced changes o ege a ion, g azing p essu e, e c., c ea es a ea-speci ic di e ences in oden abun- dance. Pa chiness will also be es ed o acknowledge in which a ea each g oup is mo e o less clumped, ega ding hei abundance alues. Al hough he exac e ec o a ea on oden abundance is no ully p edic able [ 37 ] (gi en he dis u bance g adien ) we expec ed he communal lands o ha e he lowes alues o abundance and highes pa chiness (i.e., mo e clumped), ollowed by mixed a ms and he game ese e, wi h highe abundances and lowe pa chiness; (ii) A ege a ion-s uc u e hypo hesis, i.e., a eas wi h highe he baceous co e will ha e a posi i e in luence on bo h oden size-based g oups, since i shapes he abili y o he landscape o p o ide p o ec ion agains po en ial p eda o s [21–23,25,27,38]; (iii) An ungula e-p essu e hypo hesis, i.e., oden species abundance is nega i ely in lu- enced by he abundance o ungula es, since highe g azing p essu e ends o dec ease he baceous land co e , inc ease dis u bance due o he ampling e ec , and inc ease landscape agmen a ion [9,24]; (i ) A human-dis u bance hypo hesis, i.e., oden species’ dis ibu ion is nega i ely in lu- enced by human dis u bance ac o s, such as he p esence o domes ic animals and households ha may cons ain species’ p esence [14,39]. 2. Ma e ials and Me hods 2.1. S udy A ea This s udy was implemen ed in he Mapu aland–Pondoland–Albany Biodi e si y Ho spo [ 40 ] in no he n KwaZulu-Na al, Sou h A ica. Ou speci ic s udy a ea is cha - Animals 2021,11, 2618 4 o 18 ac e ized by a spa ial g adien o human in e en ion, anging om he Mun-ya-wana p i a e game ese e (less subjec o human associa ed ac i i ies), o mixed game a ms and o communally managed lands, whe e wo dis inc Zulu communi ies a e se led (Figu e 1b). The Mun-ya-wana p i a e game ese e (27 ◦ 40 0 S–27 ◦ 55 0 S; 31 ◦ 12 0 E–32 ◦ 26 0 E) ep esen s he union o se e al p ope ies wi hou in e nal ences, managed by p i a e owne s whose goal is o explo e eco- ou is ic p oduc s, he e o e p omo ing wildli e and habi a conse a ion. Those managemen objec i es a e commonly ela ed wi h a mo e sus ainable use o wildli e, ypically wildli e- iewing ou ism [ 41 ]. The ese e is su - ounded, o he Sou h, by a mosaic o comme cial game anches o he p oduc ion o wild ungula e species, occasionally mixed wi h domes ic ca le [ 42 ] (he ea e mixed a ms) and ep esen s la ge expanses o na u al habi a wi h low human densi y. Communal lands o he eas a e composed o households, in e spe sed wi h pas u e a eas and semi-na u al ege a ion. The egion is cha ac e ized by a wa m- empe a u e clima e, wi h a humid and ho summe (Oc obe o Ap il), acco ding o he Köppen–Geige classi ica ion. Mean mon hly empe a u es ange om 19 ◦ C in July o 31 ◦ C in Janua y, and he a e age annual p ecipi a ion is 800 mm [ 43 , 44 ]. Ele a ion anges om 3 m o 304 m abo e sea le el [ 45 ], domina ed by a simila mix u e o ege a ion h oughou he a ea (bush eld, woodland and g assland) [ 46 ] (Figu e 1b). Ne e heless, he game ese e hos s a highe di e si y and abundance o p is ine habi a s, such as indigenous o es s, while mixed a ms a e mainly composed o pas u e a eas (low sh ubland and g assland–Figu e 1). Con a ily, communal lands ha e he lowes p opo ion o ege a ion and he highes co e o u ban– illage occupa ion (Figu e 1). 2.2. Roden Sampling Roden s we e sampled be ween Oc obe and No embe 2017 ( he sou he n hemi- sphe e’s sp ing) using ink- acking unnels [ 42 ], le ac i e in he ield o ou consecu i e nigh s (open ci cles in Figu e 1c). Ink- acking unnels we e made o obus co uga ed plas ic (55 × 10 × 10 cm), open on bo h ends o allow oden s o en e . Bo h en ances o he unnel a e equipped wi h an adhesi e pape wi h he glue side up, and an ink pad ( 12 ×10 cm ) was placed in he loo cen e [ 47 ] (Figu e S1B). In he middle o he unnel, a small PVC-pipe sec ion, hanging om he ceiling, was ins alled, and con ained bai composed o a mix u e o peanu bu e , oa meal and sun lowe oil [ 46 ]. The pipe was used o p e en he consump ion o he bai by he animals en e ing/c ossing he unnel. The ink unnels we e placed on he g ound, g ouped in clus e s o nine, in a Y o ma ion, 10 m apa om each o he (Figu e 1c). The a ms o he Y o ma ion we e 120 deg ees apa ( Figu e 1c ). This design p o ided an adequa e spa ial co e age in ela ion o he home- anges o he oden species, also ensu ing some le el o independence be ween sampling uni s, conside ing he mean dis ance be ween si es (see below). A e he ou - day sampling pe iod, he pla es o each ink unnel (con aining oo p in s and acks) we e pho og aphed indi idually, always a he same dis ance and wi h a e e ence scale. The oo p in da a was used o es ima e oden ela i e abundance, using he p opo - ion o he unnels wi h eco ds ( ack index; TI– o mo e de ails see Supplemen a y Ma e i- als) [ 48 ]. To ensu e ha his app oach cap u ed spa ial he e ogenei y in ela i e abundance, we conduc ed a small ial, compa ing he abundance indices de i ed om ink unnels o hose ob ained om li e- apping (see Supplemen a y Ma e ials, PART A). As ack iden i ica ion a he species le el is e y ime consuming and no iable in la ge-scale s udies, and as dis inguishing oo p in s om simila -sized species is e y di icul and bias p one, we op ed o di iding acks in o g oups based on ack size ( o mo e de ails see Supplemen a y Ma e ials PART A; Figu es S1A and S2A, Table S1A). Roden oo p in s we e g ouped in o h ee di e en size-based g oups pe body leng h/weigh , assuming a ela ion be ween oden body leng h/weigh and oo p in sizes [ 49 , 50 ]: small (body leng h: 50–100 mm), medium (100–150 mm) and la ge oden s (150–200 mm) ( Figu e S2B ). Sampling in en ionally ook place ou side he b eeding season (which peaks in he we season, [ 51 ]), in o de o a oid g ouping ju eniles in he w ong size-based g oup. Howe e , Animals 2021,11, 2618 5 o 18 conside ing he low numbe o de ec ions o la ge oden s in ink- acking unnels, we only analyzed he da a om small- and medium-sized oden s (see Resul s). The mos common species cap u ed du ing li e apping and linked o each g oup we e Mus minu oides and Dend omus melano is o small oden s, Mas omys na alensis and Saccos omus campes is o medium oden s and O omys angoniensis and Ra us a us o la ge oden s (Table S2A). Animals 2021, 11, x 5 o 18 2. Ma e ials and Me hods 2.1. S udy A ea This s udy was implemen ed in he Mapu aland–Pondoland–Albany Biodi e si y Ho spo [40] in no he n KwaZulu-Na al, Sou h A ica. Ou speci ic s udy a ea is cha ac- e ized by a spa ial g adien o human in e en ion, anging om he Mun-ya-wana p i- a e game ese e (less subjec o human associa ed ac i i ies), o mixed game a ms and o communally managed lands, whe e wo dis inc Zulu communi ies a e se led (Figu e 1b). The Mun-ya-wana p i a e game ese e (27°40′ S–27°55′ S; 31°12′ E–32°26′ E) ep e- sen s he union o se e al p ope ies wi hou in e nal ences, managed by p i a e owne s whose goal is o explo e eco- ou is ic p oduc s, he e o e p omo ing wildli e and habi a conse a ion. Those managemen objec i es a e commonly ela ed wi h a mo e sus aina- ble use o wildli e, ypically wildli e- iewing ou ism [41]. The ese e is su ounded, o he Sou h, by a mosaic o comme cial game anches o he p oduc ion o wild ungula e species, occasionally mixed wi h domes ic ca le [42] (he ea e mixed a ms) and ep e- sen s la ge expanses o na u al habi a wi h low human densi y. Communal lands o he eas a e composed o households, in e spe sed wi h pas u e a eas and semi-na u al eg- e a ion. The egion is cha ac e ized by a wa m- empe a u e clima e, wi h a humid and ho summe (Oc obe o Ap il), acco ding o he Köppen–Geige classi ica ion. Mean mon hly empe a u es ange om 19 °C in July o 31 °C in Janua y, and he a e age annual p ecip- i a ion is 800 mm [43,44]. Ele a ion anges om 3 m o 304 m abo e sea le el [45], domi- na ed by a simila mix u e o ege a ion h oughou he a ea (bush eld, woodland and g assland) [46] (Figu e 1b). Ne e heless, he game ese e hos s a highe di e si y and abundance o p is ine habi a s , such as indigenous o es s, while mixed a ms a e mainly composed o pas u e a eas (low sh ubland and g assland–Figu e 1). Con a ily, commu- nal lands ha e he lowes p opo ion o ege a ion and he highes co e o u ban– illage occupa ion (Figu e 1). Figu e 1. Loca ion o he s udy a ea in Sou h A ica, wi h he black do ep esen ing he loca ion o he s udy a ea in he Mapu aland egion o no he n KwaZulu-Na al ( a ); landscape composi ion o he h ee s udied a eas wi h dis inc managemen schemes–Mun-ya-wana p i a e game ese e, mixed a ms and communal land (Zulu ibal land)–wi h he loca ion o he sampling poin s and he numbe o sampling poin s pe a ea (in pa en hesis) ( b ); each sampling poin included a came a ap in he cen e and nine ink unnels, dis ibu ed in a Y shape (open ci cles ep esen ink unnels) (c). 2.3. En i onmen al Va iables Vege a ion s uc u e a iables we e collec ed using wo di e en app oaches: ield measu es and emo e-sensed p oduc s [ 52 ]. All a iables collec ed ha e been p e iously de ec ed as in luen ial o oden p esence elsewhe e (e.g., ege a ion co e ) [ 21 , 23 ]. Sh ub- and-g ass co e we e isually es ima ed and assigned he co esponding Edwa ds classi i- ca ion ca ego y [ 53 ] (see Table 1 o de ails), wi hin a 30 m adius bu e , cen e ed on he ink unnel’s Y o ma ion. Rega ding he land use, he p edominan ca ego ies we e selec ed ( hicke , g assland, sand o es and u ban illages) and, o each bu e , was assigned he ca ego y wi h he highes co e . Acco ding o he ype o c ops p esen in he s udy a ea, he ha es ing season occu s mos ly be ween Ap il and June [ 54 ], no coinciding wi h he s udy pe iod. The e o e, we assumed ha he e would be no in luence o c op p oduc i i y on he dis ibu ion/abundance o oden s in ou s udy. The pe cen age o ee co e was assessed based on he Global Fo es Wa ch da abase (Table 1). We also selec ed he No malized Di e ence Vege a ion Index (NDVI), widely used as a ege a ion p oduc i i y p oxy, collec ed om Landsa 8 Images [55]. Animals 2021,11, 2618 6 o 18 Va iables o ungula e p essu e and human dis u bance we e collec ed om Cu ei a- San os e al. [ 12 ] came a- ap su eys. Came as, loca ed in he cen e o he Y o ma ion, we e ac i e o 60–90 days, and a ached o a ee o me al s ake, 30 cm abo e he g ound, wi hou any bai and se o pho og aph a minimum delay (1 s o day ime and 30 s o nigh - ime) (see [ 12 ] o de ails). Each o he de ined ink- unnel clus e s (i.e., one clus e includes nine ink unnels and one came a- ap; Figu e 1c) we e spaced app oxima ely 1.4 km apa (Figu e 1b). In o al, we e sampled 196 poin s: 100 poin s in Mun-ya-wana eco- ou ism/game ese e, 50 poin s in mixed a ms and 46 poin s in communal lands. Cap u e a es, exp essed as he numbe o independen came a eco ds (>1 h in e al be ween pho og aphs o he same species, pe 100 ap-days) o li es ock (cows and goa s), wild ungula es and human dis u bance, we e used as su oga es o dis u bance in he modeling p ocedu e (Table 1). Wild ungula es we e g ouped acco ding o wo c i e ia: weigh , since ampling is one o he main nega i e impac s o ungula es on oden s [ 26 ], and/o he ac ha hey a e ac i ely managed in all s udied a eas (Table S1B). Only ungula es weighing be ween 45–200 kg and ac i ely managed we e used in he analysis, since hey a e mo e abundan han o he ungula es, as hey a e p esen h oughou he a eas unde s udy, and because hey ha e a g ea e impac on oden s, due o hei weigh (Table S1B). Li es ock we e also sepa a ed in wo weigh classes: i.e., goa s and cows. Table 1. En i onmen al a iables used in he modeling p ocedu e used o assess he de e minan s o oden abundance, collec ed in he ield, om came a- apping o based on emo e-sensing da a (GIS-based a iables). The a iable desc ip ion, ac onym, ange, esolu ion and sou ce, as well he e e ence ha suppo hei in luence on oden p esence/abundance, a e lis ed. H1—Hypo hesis 1; H2—Hypo hesis 2; H3—Hypo hesis 3, H4—Hypo hesis 4. Va iable Ac onym Desc ip ion Mean/Range Resolu ion Sou ce Suppo ing Re e ences AREA TYPE (H1) A ea Managmen con ex Mixed a ms Mun-ya-wana Communal lands Collec ed a poin - [37] VEGETATION STUCTURE (H2) T ee_Co e % T ee Co e 30.80/6–72% 30 ×30 m Global Fo es Wa ch h ps://www. global o es wa ch.o g/ (16 Ap il 2019) [27,56] Sh ub_Co e % o Sh ub co e Con inuous (C)—76–100% Semi-con inuous (SC)—51–74% Mode a ed closed (MC)—26–50% Semi-open (SO)—11–25% Open (O)—0–10% 30 m bu e Visually es ima ed [22,27,39,57–60] G ass_Co e % o G ass co e Con inuous (C)—76–100% Semi-con inuous (SC)—51–74% Mode a ed closed (MC)—26–50% Semi-open (SO)—11–25% Open (O)—0–10% 30 m bu e Visually es ima ed [22,25,38,57] Land_use Land use ca ego ies Thicke G assland Sand Fo es U ban Villages 30 m bu e 2013–2014 Na ional Land Co e Sou h A ica-SASDI h p://www.sasdi.ne / (16 Ap il 2019) [21–23] NDVI No malized di e ence ege a ion index calcula ed om Landsa images 0.48/0.28–0.67 30 ×30 m Landsa 8 h ps: //ea hexplo e .usgs.go / (18 Ap il 2019) [60,61] Animals 2021,11, 2618 7 o 18 Table 1. Con . Va iable Ac onym Desc ip ion Mean/Range Resolu ion Sou ce Suppo ing Re e ences UNGULATE PRESSURE (H3) Goa s Cap u e a e o goa s (numbe o eco ds pe 100 days o apping) 0.16/0–1.88 Collec ed a poin Came a- apping su ey [9,24,26] Li es ock Cap u e a e o cows (numbe o eco ds pe 100 days o apping) 0.20/0–3.17 Collec ed a poin Came a- apping su ey Wild Ungula es Cap u e a e o ungula es (numbe o eco ds pe 100 days o apping) 0.750/0–3.48 Collec ed a poin Came a- apping su ey DISTURBANCE VARIABLES (H4) HUMANS Cap u e a e o humans 0.84/0–10 Collec ed a poin Came a- apping su ey [39] DIST Dis ance o houses 2.738/0.031–9.867 km Collec ed a poin Came a- apping su ey 2.4. Da a Analyses/Modelling 2.4.1. Spa ial Pa e ns o Roden Rela i e Abundance Ac oss A eas and Size-Based G oups Di e ences in mean abundance alues o size-based g oups (small and medium) be ween s udy a eas (Mun-ya-wana game ese e, mixed a ms and communal lands) we e es ed using GLM wi h 3-le el a ea co a ia e and binomial e o dis ibu ion. The magni- ude o pa chiness in each a ea was asce ained by spa ial-poin pa e n analysis o coun da a using Lloyd’s index o pa chiness [ 62 ]. A Lloyd’s index o 1 indica es a andom dis ibu ion, whils one <1 sugges s uni o mi y and >1 pa chiness. 2.4.2. In luence o En i onmen al Va iables on Roden Rela i e Abundance Due o he high numbe o candida e a iables and o a oid mul icollinea i y bias, we i s es ima ed he nonpa ame ic Spea man’s co ela ion ( s ) using he “psych” R package [ 63 ]. When a high co ela ion be ween wo co a ia es was de ec ed ( s≥ 0.7; [ 62 ]), he a iable ha was less co ela ed wi h he dependen a iable was excluded om he analysis [64]. The in luence o all candida e a iables on oden ela i e abundance was es ed using a boos ed- eg ession- ee (BRT) app oach, implemen ed wi h he “gbm” package [ 65 ] in R [ 66 , 67 ]. This modelling echnique encompasses he ad an ages o eg ession ees (e.g., p edic o a iables can be o any ype, analysis is insensi i e o ou lie s and can accommoda e missing da a [ 68 ]), o e coming hei low p edic i e capaci y h ough he boos ing algo i hm [ 69 ]. The inal model is a linea addi ion o se e al eg ession models in which he simples e m is a ee [68,70]. Boos ed- eg ession- ee models a e esilien o model o e i ing bu , o ha e a be - e p edic i e pe o mance, we de ined, a p io i, he model’s inpu pa ame e s based on Ca slaw and Taylo ’s sugges ions [ 70 ]. In BRT, lea ning a e (l ) is he sh inkage pa ame e ha con ols he con ibu ion o each ee o he model, and ee complexi y ( c) de e mines he numbe o nodes in a ee and, consequen ly, i s size. These wo pa ame e s con ol he numbe o ees in he model, while he bag ac ion (0.5) selec s he p opo ion o da a be- ing used a each s ep [ 61 , 70 , 71 ]. All models we e i ed o allow in e ac ions using a en- old c oss alida ion o de e mine he op imal numbe o ees o each model. The la ges lea n- ing a e and he smalles ee complexi y we e selec ed o allow a minimum o 1000 ees in he BRT i ing p ocess (see [ 68 ]). Non-in o ma i e a iables we e emo ed du ing he i ing p ocess, allowing he simpli ica ion o he se o a iables [ 68 ]. This simpli ica ion consis ed o de ining how many a iables he unc ion can es o emo e, based on ela i e Animals 2021,11, 2618 8 o 18 in luence and o al numbe o a iables. Then, a g aph was p oduced showing di e ences in he p edic ed de iance acco ding o se e al scena ios, each one wi h a di e en numbe o a iables emo ed. Nex , he numbe o a iables o elimina e was decided, and hey we e emo ed in o de o mino ela i e in luence. We de ined a h eshold alue and only epo ed he in e ac ions wi h ela i e in luence alues >10%. The inal ela i e in luence o each a iable was calcula ed by a e aging he numbe o imes a co a ia e is used o spli ing, weigh ed by he squa ed imp o emen o he model as he esul o each spli . I is hen scaled, such ha he alues sum o 100 [ 72 ]. Fi ed alues we e plo ed in ela ion o he mos impo an p edic o s, e ealing hei e ec s on oden abundance. Explained de iance was calcula ed using he ollowing o mula om Abea e (2009) [73] D2=1− esidual de iance o al de iance  The 95% con idence in e als o each a iable we e es ima ed o he i ed unc ion by aking 500 boo s ap samples o he inpu da a, wi h he same size as he o iginal da a. A BRT was i ed o each sample, and he 5 h and 95 h pe cen iles we e calcula ed o he poin s o each unc ion. Models we e buil sepa a ely o small- and medium-sized oden s. Fo each model pe o med, in e ac ions be ween ypology and he o he in luen ial independen a iables (i.e., ela i e impo ance abo e >10%) we e es ima ed, o e alua e con ex -dependency in he in luence in he e ec en i onmen al a iable associa ed wi h he managemen con ex . All analyses we e implemen ed in R ia R S udio Ve sion 1.1.463 [66,67]. 3. Resul s 3.1. Spa ial Pa e ns o Roden Abundance Ac oss A eas and Size-Based G oups F om he 192 sampling poin s moni o ed, 85% p esen ed small oden acks, while 76% de ec ed he occu ence o medium oden s, wi h an o e lap in 35% o si es and in e -a ea a ia ion in de ec ion (i.e., numbe o unnels wi h signs/ o al numbe o unnels, Table S2B). Mean abundance in Mun-ya-wana game ese e was 0.52 ± 0.26 (mean ± SD) o small oden s and 0.43 ± 0.34 o medium oden s; in mixed a ms, 0.31 ± 0.21 o small oden s and 0.52 ± 0.32 o medium oden s; and in communal lands was 0.26 ± 0.23 o small oden s and 0.36 ± 0.24 o medium oden s (Figu e 2). Rega ding he GLM esul o size-based g oups, i e ealed signi ican di e ences in ela i e abundances only o small oden s, be ween Mun-ya-wana game ese e and he emaining a eas (Table S3B, Supplemen a y Ma e ials). No signi ican di e ences we e de ec ed in ela i e abundances o medium oden s be ween a eas (Figu e 3). Be ween g oups, signi ican di e ences we e only ound in mixed a ms (Table S3B, Supplemen a y Ma e ials), wi h medium oden s being mo e abundan (0.52 ± 0.37) han small-size oden s (0.31 ± 0.26) (Figu e 3). Based on hese esul s, he e ec o en i onmen al d i e s on oden abundance was e alua ed sepa a ely o each o he size-based g oups. Roden Pa chiness Lloyd’s Index o Pa chiness e ealed ha o e e y a ea and size-based g oup, all abun- dance alues we e agg ega ed ( γ > 1; Table 2). Bo h medium and small oden s a e he e oge- neously dis ibu ed wi hin he h ee s udy a eas (Figu e 3), demons a ing a he e ogenei y g adien . Acco ding o Table 2, we can obse e ha he highes alues o small oden s a e in communal lands, ollowed by mixed a ms and inally, he game ese e. Fo medium oden s, he e is a g ea e clus e ing pa e n in he game ese e, ollowed by communal lands and mixed a ms. Wi h hese esul s, i is possible o s a e ha he abundance pa e ns di e be ween he size-based g oups, and wi hin each a ea. Animals 2021,11, 2618 9 o 18 Animals 2021, 11, x 10 o 18 Figu e 2. Map o he s udy a ea showing oden dis ibu ions: small-size oden s a e in o ange and medium-size oden s in yellow. The size o each poin is equi alen o abundance alue, as indica ed in he espec i e legend. Figu e 3. Boxplo o medium and small oden s’ ela i e abundance in he h ee managemen - ype zones moni o ed: game (mixed) a ms, Mun-ya-wana game ese e and communal lands. Based on he GLM es , * indica es a signi ican di e ence be ween size-based g oups in mixed a ms (p = 0.011), + indica es a signi ican di e ence be ween Mun-ya-wana game ese e and emaining a eas o small oden s (p = 0.016). Figu e 2. Map o he s udy a ea showing oden dis ibu ions: small-size oden s a e in o ange and medium-size oden s in yellow. The size o each poin is equi alen o abundance alue, as indica ed in he espec i e legend. Animals 2021, 11, x 10 o 18 Figu e 2. Map o he s udy a ea showing oden dis ibu ions: small-size oden s a e in o ange and medium-size oden s in yellow. The size o each poin is equi alen o abundance alue, as indica ed in he espec i e legend. Figu e 3. Boxplo o medium and small oden s’ ela i e abundance in he h ee managemen - ype zones moni o ed: game (mixed) a ms, Mun-ya-wana game ese e and communal lands. Based on he GLM es , * indica es a signi ican di e ence be ween size-based g oups in mixed a ms (p = 0.011), + indica es a signi ican di e ence be ween Mun-ya-wana game ese e and emaining a eas o small oden s (p = 0.016). Figu e 3. Boxplo o medium and small oden s’ ela i e abundance in he h ee managemen - ype zones moni o ed: game (mixed) a ms, Mun-ya-wana game ese e and communal lands. Based on he GLM es , * indica es a signi ican di e ence be ween size-based g oups in mixed a ms ( p= 0.011 ), + indica es a signi ican di e ence be ween Mun-ya-wana game ese e and emaining a eas o small oden s (p= 0.016). Animals 2021,11, 2618 16 o 18 22. Layme, V.M.G.; Lima, A.P.; Magnusson, W.E. E ec s o i e, ood a ailabili y and ege a ion on he dis ibu ion o he oden Bolomys lasiu us in an Amazonian sa anna. J. T op. Ecol. 2004,20, 183–187. [C ossRe ] 23. Holland, G.J.; Benne , A.F. Di e ing esponses o landscape change: Implica ions o small mammal assemblages in o es agmen s. Biodi e s. 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